Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. emnn ouainuisfrmngmn ln nue h rsraino vltoayptnili en- De- in diversity. potential within-species evolutionary of of preservation determination the the ensures is plans efforts management conservation for current units of population challenges termining threatened (Woinarski hundred the concern one of reached over conservation marsupials for One particular 2018). underway of al., a are proportion et them efforts high (Legge conservation making A priority mammals century, high 2015). Australian last Consequently, al., the 2011). et al., in Ziembicki et vulnerability 2015; 2018; al., of Dickman, 2017), et levels & al., Woinarski (Baker high et marsupials 2014; Crooks Australian al., 2019; in et al., precipitous et particularly Fisher (Bowyer been globally have issue declines urgent population an but is diversity mammalian of conservation The INTRODUCTION conclude genetics-based We to contribution limited. However, phenomic exceedingly rigorous seems exists. statistically diets) comprehensive, none local a because provide studies. to adaptation conservation can specialization musculoskeletal) morphometrics (e.g. geometric general variation populations 3D probably between morphological that to (and cross-breeding heritable due that skull for possibly upper indicates variation, potential morphological adaptation the shape the local local for as of reduce scope well lack within-species not as shape The uniform of muscles, will a constraints. majority masticatory suggest vast allometric the results of or Strikingly, Our areas plasticity nothing. unexplained. phenotypic insertion or remained little in - contributed changes variables 76% explained geographic - to differences and variation relating population Climatic Size-adjusted sizes, region. incisor effect shape. and cheekbone smaller muzzle the inhabited, best, far variation shape in area with skull differences shape variation predicted land and Size less skull of variation variation. size to Regardless shape skull braincase latitude/longitude overlapping with broadly analyses. and of coinciding phenotypic partitioning amounts temperature, similar variation and harbour rainfall, islands. and populations behaviour, several ANOVA, quoll sex, northern and feeding, Procrustes size, populations Components, mainland of population, four Principal proxy of the through across a changing contribution 5,000 to individuals geometric is adaptability the its 101 indicate – variation in across assessed shape of shape We quoll, sciences patterns Skull skull if northern evolutionary and analysed distinguished We the the coast. be conditions. marsupial, can environmental Australian in populations endangered if northern used an ask to of the us widely allowing skull differentiation, along technique the linear range a in taxonomy-focused diversity distribution use beyond investigated shape km we assessed characterize generally Here, rarely to is is – species diversity morphometrics descriptions. mammalian phenotypic qualitative threatened intraspecific or of However, measurements populations genetics. among population diversity through of distribution biogeographical The Abstract 2020 5, May 4 3 2 1 Guillerme populations Viacava fragmented Pietro between adaptation reveals local marsupial of endangered evidence widely-distributed, little a of shape Skull lnesUniversity Flinders Technology of University Queensland England New of University Queensland of University The 1 keCameron Skye , 1 ioeBlomberg Simone , 1 .S Wilson S. R. , 1 areeSansalone Gabriele , 1 n eaWeisbecker Vera and , 1 2 ate Phillips Matthew , 4 3 Thomas , Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. eas hyihbtawd ag fhbtt,rnigfo anoet odsrs(eg 91 or tal., carrion et and Moore fruit invertebrates, 1981; vertebrates, (Begg, investigation small deserts this of to for foragers rainforests system from opportunistic island study ranging are several habitats, They suitable and of 2002). a 2010) range Oakwood, Ward, also information wide 2019; and are a no biogeographic (Hill quolls inhabit but flow major Northern they gene by 2015) km 1999). because apparent separated al., 5000 al., no now over et et with (Woinarski are distribution Woolley populations populations They pre-colonial 2009; mainland 1994). a al., four Griffiths, had into et & have breaks How (Braithwaite to Australia 2016; appear al., northern al., quolls et across et Firestone Northern on 2009; adaptation. Hohnen focus al., morphological 2010; We et on Ward, concern. 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(Kendall, size) minus (form 4 Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. 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(Table 0.001) < p 0.121, = R-squared 3.125, = (1,99) 41,Rsurd=015 < p 0.125, = R-squared 14.15, = (1,90) 39 R- 23.9, = Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. oipoeorudrtnigo hte h pce ersn opooia rfntoa conservation was functional aim or Our morphological divergences represent range. genetic species longitudinal and the 5000-kilometre adaptive whether their biogeographic on across the understanding evident that our be expectation improve the should to was quolls study northern the this between represent of not rationale do The and schematic unexplained are of sizes amount Circle The model. (76%). DISCUSSION the shown. residuals by not are the explained and values by negligible shape individual inner depicted is of The the is [g] amount and variables. model variables explanatory [c] this three the corresponding climate by all are the pure of numbers shape of interaction and outer interaction the [b] The the for size set. of fraction pure each values shape. 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Slopes of Table of Association Homogeneity on of shown in Results are 2008). role populations Klingenberg, of a & (Drake slopes play size on not shape of does regression allometry 3 Thus Figure populations. populations. between of popu- shape slope differences between the allometric significant differences differentiating populations; same similar significant between revealed the show differences 0.203) allometry following shapes shape = of populations the (p residuals reveals of (F the population also lations hypothesis on each shape analysis the performed Homogeneity of of shape ANOVA A that determinant slopes Allometry-corrected Procrustes variation. meaning allometric strongest rejected. shape 3), among the total not the differences Figure as is of significant and manifests % no 1 15.1 size therefore for suggests (Table tested, We accounting Test variables 0.001), analyses. 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Data may be preliminary. u hsi ueybhvorlcneuneo h uln cino h eprlsmsl (Washburn, muscle temporalis the of action pulling 2006), al., the et of (Flores It age consequence individuals. with quoll crests behavioural the northern sagittal Perhaps purely larger among display skull. length a to in the is – widely of males varies this uses the particularly which but individual – in crest, variation mammals on sagittal for the based the common of is process is much this re-modelling that of a possible example of from also best predictor derives is shape. it main skull in adaptation, quoll a of variation microevolutionary northern lack is little of the precipitation explains hypotheses the variable as although from to this well Aside addition, more 2015), due as In of al., doubtful populations, et consisting is between shape. 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(Wroe Eylandt, or within bite vertebrates Territory Groote habit greater Northern or as imply feeding such Pilbara to environments, or therefore as wetter could is diet to such populations It compared particular some environments, when between a 2019). drier placement al., to in arch et adaptation zygomatic Weisbecker populations and as 2019; length popula- Wroe, interpreted muzzle quoll 2011; & in be (Meloro, northern Mitchell variation mastication the some 2018; in that distinguish al., changes possible to et to adaptation Sansalone, appear evolutionary well-known Sherratt, muzzle arches. a Mitchell, the zygomatic mirrors variation and wider anteri- This arch and more tions. nasals sexes zygomatic a shorter between the crest, braincases, Differences sagittal of smaller skulls larger Rearrangements arch. crests, a larger zygomatic sagittal skull, dorsally-oriented larger the variation: with associated of of interpreted. males rest and regions visually include the scar to main be masseteric relative two can positioned braincases identifies orly sex smaller overall allometry and have by population, to predicted tend size, displace- displacement to the course quolls, landmark according northern of The the configurations traits. although among physiological) landmark divergence adaptation, or shape between local of (behavioural ment implications differential Kelly non-morphological functional on to the 2009; understand based due al., better This investigated To effect et further adverse (Cardoso genetic be differentiations. any populations to on morphological risk need outbreeding adaptive based would not Thus, of this would with size benefits hypothesis. 2019) coincide of fitness constraints Phillips, genetic to independently developmental & that appear vary the indication not to contradicting an do again shape provides divergences population, skull population or of sex the ability are possibly to stochastic, the sex adaptation. related a local and demonstrate reflect suggest factors to populations also might appear This sizes, not effects does account. effect however, These marsupials which, into low populations taken between other had divergence is shallow in size populations heritable, when between findings even shape to distinguishable, ways several skull statistically contrary of intriguing of one is an differentiation represent species. might This is Although the and it within 2019), However, al., level. traits of among et morphological Weisbecker within-species size overlap strong. shape 2018; and broad the to al., be shape et the skull at to Sansalone, between and Sherratt, unlikely visible association explains, (Mitchell, evolutionary is be size larger-scale constraint differentiate the may that to of such (~16%) marsupials some seem any variation least populations at shape, of the that and amount and indication variation low size shape adaptability the in the most given reduce populations the However, Queensland might explains constraints eastern size developmental size. as the that by skulls, concern of quoll our length for seems northern western the support of also localized limited across It more some individuals is that sex. the as There such different of shape of populations, in ends are the they disparate of opposite each if as seaboard. from even within just individuals distributed shape, appear sizes evenly in similarly-sized populations effect is similar that low variation be have meaning most to differences that size, likely sex are than shape and distribution of variation populations discrete, biogeographic (~76% into particular, less evolved variation have In explain shape populations the morphotypes. and quoll the of adapted, any northern of that management locally in evidence c 7 Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. eg .J 18) h ml aml fLtl orageRc,NTII clg of Ecology III. N.T Rock, Nourlangie Little of mammals small The https://doi.org/10.1071/wr9810073 antechinus, (1981). swamp hallucatus the J. rus of R. redescription Begg, and Taxonomy (2015). 127–170. S. Dyck, Van minimus & dusky ( A., Australian antechinus Baker, the dusky of Museum–Nature mainland Queensland assessment the the taxonomic of of A ( species antechinus Memoirs to (2015). dusky Peninsula elevation L. Tasman an E. the Gray, and species, & new A E., genomes. Mason, complex: to antechinus T., Allozymes Mutton, A., populations: Baker, (2018). natural C. of Dickman, of & conservation A., analysis the Baker, and and collection Genetics the for Ecology (2017). package Molecular R W. An F. geomorph: Allendorf, (2013). data. E. shape Otarola-Castillo, morphometric geometric & of C., analysis D. The Adams, Formation: Chanares the of cynodonts Gomphodont 4634(2000)020[0501:GCOTCA]2.0.CO;2 (2000). P. sequence. ontogenetic N. an Giannini, & F., of Abdala, capacity intrinsic less show REFERENCES marsupials with iv. if comparison identify mammals. a to addition, placental and useful in than biomechanical clade; be variation this threatened adding would shape in other species and exists in placental predictors, that variation variation similar as marsupial the shape ecologically time dissect patterns of that further short or scope is to within variables the studies adaptations suggestion developmental identifying climatic specific concerning into other evolve directed the suggested investigating not be together, been but marsupials, plasticity, should also kangaroos; research individual similar Further has as for A well apparatus scope spans. species. skull a masticatory as have the quoll the species might within northern wombat in mammals variation of plasticity living heritable determination larger-scale, within-species the for the apparatus high for the scope masticatory in adaptability potentially across little the mechanism low of differentiation in is a response be pattern variation of there suggests major might shape that lack it of a suggesting a there the because concentration shape, is where hand, if concern The plasticity point a other investigated change. individual is environmental the the be suggests to populations On to to species or adapted the needs conditions, so). of locally climatic it do biomes, not (although to of are reasons indicated diversity populations physiological is different or phenomes low behavioural of of implications. population important individuals find of two overarching that has The separation variation, suggests marsupial. unexplained endangered of it levels an First, data high of phenomic very distribution quantitative, and the useful, differentiation, adds across morphological shape variation skull of populations. quoll assessments northern between to of genetic 2019) analyses as al., morphometric decisive such bite et of geometric a factors (Weisbecker higher The tendency not non-adaptive feeding to are potentially weak in related populations from the variation between originate are individual with differences rather muzzles or the consistent and shorter that drift survival, also as suggests individuals’ is size, again the This to of This (Braithwaite factor regardless 2007). bite” males. Milne, females, & “mating larger than (Wroe PC1 muzzles the the forces along shorter possibly of found have and apparatus, 2000) to behaviours masticatory Oakwood, males the masticatory 1994; on from variation 2019). Griffiths, arise al., shape & pattern, et zygomatic of kangaroos Weisbecker allometric emphasis and 2018; the wombats the the al., for in et that and suspected Sansalone, suggested possible also Sherratt, therefore been Mitchell, is is 2018; and has It al., 1991), lifetime Ravosa, et Ledogar, 2000; individual’s Sherratt, Giannini, an (Mitchell, & (Abdala during mammals re-modelling of arch bone Similarly, 1947). E effo)(aspai:Dasyuridae). (Marsupialia: Geoffroy) (E. h otenqol(aspai:Dasyuridae). (Marsupialia: quoll northern the , , 26 2,4040 https://doi.org/10.1111/mec.13948 420–430. (2), ora fVrert Paleontology Vertebrate of Journal ertlvso anvru marsupials carnivorous of lives Secret ehd nEooyadEvolution and Ecology in Methods , 59 75–126. , 8 eor fteQenln uem-Nature - Museum Queensland the of Memoirs , 20 3,5156 https://doi.org/10.1671/0272- 501–506. (3), nehnssanoi mimetes swainsonii Antechinus idieResearch Wildlife , 4 SR PUBLISHING. CSIRO . 4,393–399. (4), nehnsvandycki Antechinus , 8 1,73–85. (1), (Thomas)). Antechinus p Nov.) sp. Dasyu- , 59 , Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. e ohs . ot .M,Tre,N,Bie,J . edy .P,Knio,M . cwizr .A,& A., J. Schweitzer, T., M. Kinnison, variation. P., intraspecific A. of Hendry, importance K., ecological J. Consider- Bailey, The 2 L. N., Turley, (2018). Boitani, P. M., & E. D. (2000). C., Palkovacs, mammals. Post, Rondinini, S., M., K. terrestrial Roches, Marco, in Di Des R. Sciences risk B., of extinction R. Wayne, Academy reveals S. National fragmentation King, & the M., habitat of D. of M., Theobald, Quantification L., G. C. (2017). Burdett, Mace, R., K. P., Crooks, biology. R. conservation in O. https://doi.org/10.1016/S0169-5347(00)01876-0 processes Bininda-Emonds, and evolutionary skull problem. A., the ing 3D between K. co-variation a shape for Crandall, approach Does 3D (2013). A https://doi.org/10.1111/joa.12086 A. 329–336. consequences? Herrel, (4), & functional pheno- for have T., change mandible Souter, phenotypic of the M., analysis Baylac, for method R., A Cornette, (2015). C. data. D. high-dimensional Adams, by & described J., types D. adaptation? Sekora, ecophenotypic L., distinc- or M. Cranial effect Collyer, drift (2012). Genetic P. Society Linnean Ciucci, bear: Bunce, the & brown of P., A., Apennine Taglianti, Gomerˇci´c, Prentis, the Vigna D., in R., T., Huber, tiveness of A. A., systematics Loy, Evans, the P., of Colangelo, W., assessment Dodt, morphometric and C., molecular A Burg, Society (2019). Der J. Van Implica- M. B. D., Phillips, the K. & populations: Haouchar, C., Firestone, island Fruciano, M., & M., translocated Cascini, B., of M., quoll. W. variation Celik, northern Sherwin, genetic the B., the of Rankmore, on management M., events https://doi.org/10.1007/s10592-008-9774-z conservation Oakwood, founder for B., of tions D. Effects M. Eldridge, (2009). the J., in M. contraction Cardoso, range and variation . morphometrics. Geometric Demographic (2013). A. Cardini, (1994). D. (2010). A. G., A. https://doi.org/10.1071/WR9940203 A. Griffiths, E. Elkahloun, quoll, Ostrander, & A., northern W., K., . . Reynolds, . Zhao, R. J., A., dogs. E., Novembre, Braithwaite, in Auton, K. S., variation Lohmueller, G. M., morphological Johnson, D., Cargill, underlies B., J. architecture https://doi.org/10.1371/journal.pbio.1000451 M., N. Degenhardt, genetic Sutter, B. J., simple S., vonHoldt, A J. D. G., Mosher, Schoenebeck, mam- M., H. L., world’s Castelhano, Parker, the Li, of P., A., Conservation Quignon, Brisbin, R., (2019). L. A. J. Boyko, Rachlow, hunting. & and principal R., efforts, of community J. https://doi.org/10.1093/jmammal/gyy180 areas, means Goheen, protected S., by Status, M. data mals: Boyce, ecological T., of R. Bowyer, analysis spatial All-scale change. matrices. 3800(01)00501-4 (2002). landscape neighbour P. and of Legendre, fragmentation coordinates & Habitat D., (2010). Borcard, A. D. All Saunders, for & Biology F., A. Bennett, 1,5–4 https://doi.org/10.1038/s41559-017-0402-5 57–64. (1), Macropus , 186 3,7382 https://doi.org/10.1093/zoolinnean/zlz005 793–812. (3), ope lrfistetmoadmd fknao evolution. kangaroo of mode and tempo the clarifies complex , ayrshallucatus Dasyurus 93 1544–1550. , , 107 1,1–6 https://doi.org/10.1111/j.1095-8312.2012.01926.x 15–26. (1), clgclModelling Ecological , Mruila Dasyuridae). (Marsupialia: 114 Heredity 2) 6574.https://doi.org/10.1073/pnas.1705769114 7635–7640. (29), ilgclSineFnaetladSseais K Oxford UK: Systematics. and Fundamental Science Biological , 115 9 rnsi clg Evolution & Ecology in Trends , 4,3735 https://doi.org/10.1038/hdy.2014.75 357–365. (4), 153 osrainGenetics Conservation 1,5–8 https://doi.org/10.1016/S0304- 51–68. (1), ora fMammalogy of Journal idieResearch Wildlife olgclJunlo h Linnean the of Journal Zoological LSBiology PLOS aueEooy&Evolution & Ecology Nature ora fAnatomy of Journal , , , 10 , ilgclJournal Biological 15 100 21 , 8 6,1719–1733. (6), 8,e1000451. (8), 7,290–295. (7), 2,203–217. (2), Conservation 3,923–941. (3), Proceedings , 223 , Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. uz . itrekr .(03.Smlnmrs ehdfrqatfigcre n surfaces. and curves quantifying Mammalogy for of method Journal A Italian Semilandmarks: (2013). the P. Mitteroecker, & P., Gunz, Dasyurus. of (2019). https://doi.org/10.5281/zenodo.2620785 species V. Weisbecker, new & a T., and Guillerme, Perameles, of species High-density new (2019). a D. of Lond P. Characters Polly, (1842). not-really-a-problem. & J. the Gould, C., and A. bad, the Fabre, good, C., Biology The Comparative Bardua, integration: oral and and N., marsupial shape R. of the Felice, Sanchez-Villagra, analysis of morphometric A., & evolution Watanabe, L., the A., Hautier, limit Goswami, integration V., high C. and Bennett, constraints V., developmental Weisbecker, apparatus? Do D., (2016). P. integration Polly, R. and M. M., variance Randau, Shape, A., (2012). Goswami, R. mammals. M. placental Sanchez-Villagra, https://doi.org/10.1111/j.1420-9101.2012.02477.x and 862–872. & (5), marsupial O., aus- Contrasting Mock, the craniogenesis: D., in during P. skull Polly, the A., of Goswami, ontogeny postnatal Comparative (2006). F. Morphology Abdala, & metatherian N., tralidelphian Giannini, J., The VanDerWal, A., (2014). P., A. D. S. Kutt, repeating? Flores, & Blomberg, history J., Is H., I. Gordon, Australia: McCallum, A., in A., Fisher, marsupials R., S. 23 C. tropical Fritz, Thomas, of J., M., decline units Letnic, M. S., of current Lawes, Legge, differentiation conservation A., N., Frank, and C. B., Variability and Abbott, Johnson, (2000). O., of, E. D. Geffen, Fisher, Origin & B., (2008). W. genus Sherwin, M. marsupials. the A., J. B. in Houlden, microsatellites Rochford, B., K. & ( Firestone, J., squirrel the in red C. change a Irish https://doi.org/10.1007/s10592-007-9419-7 Edwards, of victim the N. a A., C. in, quoll Johnson, eastern L. & the E., Is M. Finnegan, decline: Jones, species E., in C. change Hawkins, climate study C., of weather? S. case role Nicol, A the J., VanDerWal, Testing carnivores: L., (2015). small B. in Bateman, flexibility B., Fancourt, Dietary (2017). S. quoll, T. northern Doherty, ps://doi.org/10.1093/jmammal/gyx015 endangered & the K., Rayner, on functional A., Do J. (2016). musteloid Dunlop, in C. evolution A. shape Fabre, skull drive & pattern Peign´e, activity S., A., and carnivorans? diet, Goswami, habitat, L., locomotor with Botton-Divet, associated E., demands C. of Wall, transformation M., Historical Dumont, and change: Morphological morphological of dogs. pace Bernard The (2015). https://doi.org/10.1098/rspb.2007.1169 St (2008). V. in P. C. shape S. Klingenberg, skull & Edwards, G., ( & A. conservation. Albatrosses Drake, T., for Black-footed Implications Hiraoka, Japanese F., RADseq: https://doi.org/10.1111/eva.12274 and Sato, digest Hawaiian double J., of ing A. comparisons Shultz, genomic G., E. Dierickx, 2,1110 https://doi.org/10.1111/geb.12088 181–190. (2), 41–42. , LSONE PLOS nertv n oprtv Biology Comparative and Integrative osrainGenetics Conservation , ilgclJunlo h ina Society Linnean the of Journal Biological 267 4,4640 https://doi.org/10.1002/jmor.10420 426–440. (4), , 10 , ayrsalbopunctatus Dasyurus 6,e192.https://doi.org/10.1371/journal.pone.0129420 e0129420. (6), 59 Dasyurus cuu vulgaris Sciurus 3,6963 https://doi.org/10.1093/icb/icz120 669–683. (3), rceig fteRylSceyB ilgclSciences Biological B: Society Royal the of Proceedings , ayrshallucatus Dasyurus , 24 1 n osrainipiain o h ag utaincarnivorous Australian large the for implications conservation and 2,1513 https://doi.org/10.1023/A:1026578821339 115–133. (2), 1,103–109. (1), adR ol o esrn adakpsto variation position landmark measuring for Tools LandvR: population. ) , 56 Mruila ayrmrh:Dasyuridae). Dasyuromorpha: (Marsupialia: 10 3,4445 https://doi.org/10.1093/icb/icw039 404–415. (3), , 117 . ora fMammalogy of Journal 4,8888 https://doi.org/10.1111/bij.12719 858–878. (4), osrainGenetics Conservation vltoayApplications Evolutionary ora fEouinr Biology Evolutionary of Journal lblEooyadBiogeography and Ecology Global heati nigripes Phoebastria , 98 , , 275 9 3,8886 htt- 858–866. (3), , 8 5,1099–1109. (5), rc ol Soc. Zool. Proc. 13) 71–76. (1630), 7,662–678. (7), Integrative ora of Journal Zenodo. . Hystrix, us- ) , 25 , Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. ari,G,&Ceeu,J 21) iea ieo es eitneI:Drc eeto nsz rcorrelated or size on selection Direct II: resistance least of line a as constraints? mor- Size to adaptive (2010). due response J. and Cheverud, Diet & resistance: G., evolutionary Marroig, least of line monkeys. a world as 3820.2005.tb01049.x new Size in (2005). radiation and J. phological force Cheverud, bite & Diet, G., (2016). O. Marroig, R. T. Freitas, de (2018). morphotype. & rodent B. R., https://doi.org/10.1111/jeb.12937 generalist L. Wintle, the Monteiro, in & R., morphology Fornel, D., skull D., B. Southwell, Patterson, B., R., Maestri, Scheele, D., communities ecological Lindenmayer, and N., species threatened Robinson, variation. S., genetic hidden Legge, of potential Evolutionary (2008). O. Evolution shape. marsupial. Carlborg, & of endangered & Theory A., an statistical Rouzic, the in Le of adaptation survey rapid A (1989). and G. flow D. gene Kendall, Targeted (2019). L. Biology B. for Conservation value Phillips, conservation & high process. E., have threatening Kelly, populations a Island of (2009). H. ahead L. https://doi.org/10.1111/j.1469-7998.2009.00569.x predator Schmitt, 206–217. marsupial & top S., Australia’s B. P. northern quoll Spencer, northern A., declining R. Burridge, How, the & of N., C. populations Johnson, among J., flow I. z gene Radford, S., predict B. topography P. and Spencer, ( Rainfall M., Hillyer, (2016). A S., P. Legge, Europe: C. D., in K. genetics Tuft, population R., Wolf Hohnen, (2017). management. U. and D., conservation Saarma, https://doi.org/10.1111/brv.12298 S. for . 1601–1629. Baltr¯unait´e, . suggestions Czarnomska, . (3), and V., L., J., meta-analysis Aspi, A. review, D., Stronen, systematic Rungis, quoll C., R., J., Nowak, northern Ozolins, Godinho, Vil`a, E., C., the J., Randi, M., for A., Pilot, plan J. J., Leonard, recovery Darwin Remm, Sport, National M., and (2010). Arts Hindrikson, J. The Environment, S. Resources, Ward, Natural mammals: of & in Department skull M., the B. and muscles Hill, Jaw (2001). D. C. Marshall, & & Physiology O., J., mastication. C. Z. of D., Liu, Justice, biomechanics L., Thau, L., The K. Rafferty, change. Chini, A., W., cover Tyukavina, A., S. forest Egorov, Herring, A., 21st-century A., of S. Kommareddy, maps Turubanova, R., global https://doi.org/10.1126/science.1244693 M., High-resolution T. 850–853. Hancher, (2013). Loveland, (6160), G. J., R., R. S. Moore, J. Townshend, Goetz, V., V., P. S. Potapov, Stehman, C., M. (2015). Hansen, R. L., J. Townshend, E., B. Press. (1993). Foster, T. . Chicago K. . M., Lovejoy, . of B. R., R. Hall, D., & C. Ewers, R. J., ecosystems. Margules, I., Hanken, Holt, J., Earth’s E. D. on Damschen, A., Levey, impact M., Gonzalez, F., lasting W. W. F., https://doi.org/10.1126/sciadv.1500052 its Cook, Laurance, K. and D., J., fragmentation Davies, C. A. Habitat Collins, King, J., P., N., Clobert, M. C. A., Austin, Jenkins, O., L. J. Brudvig, Sexton, M., N. Haddad, ayrshallucatus Dasyurus , 131 , 23 1,2729 https://doi.org/10.1016/S1095-6433(01)00472-X 207–219. (1), 1,3–7 https://doi.org/10.1016/j.tree.2007.09.014 33–37. (1), ). , 33 osrainGenetics Conservation 1,1211 https://doi.org/10.1111/cobi.13149 112–121. (1), Evolution oprtv iceityadPyilg atA oeua Integrative & Molecular A: Part Physiology and Biochemistry Comparative h kl,vlm :Fntoa n vltoaymechanisms evolutionary and Functional 3: volume skull, The , 64 kso,M,Loe-a,J V., L´opez-Bao, J. M., Akesson, ˚ 5,17–48 https://doi.org/10.1111/j.1558-5646.2009.00920.x 1470–1488. (5), Evolution sr Publishing. Csiro . , 17 5,11–28 https://doi.org/10.1007/s10592-016-0856- 1213–1228. (5), 11 ora fEouinr Biology Evolutionary of Journal , 59 5,12–12 https://doi.org/10.1111/j.0014- 1128–1142. (5), ttsia Science Statistical cec Advances Science lae,F,Laea . Echegaray, L., Llaneza, F., Alvares, ´ ora fZoology of Journal . ilgclReviews Biological , ayrshallucatus Dasyurus 4 , 29 2,8–9 JSTOR. 87–99. (2), , rnsi Ecology in Trends 1 1) 2191–2204. (11), 2,e1500052. (2), Science University . Monitoring , 278 , , (3), 342 92 . Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. ot,A,Sia,L . eOier,F . ari,G 21) ievrain rwhstrate- growth variation, Size (2013). G. Marroig, skull. & mammalian B., the in F. modularity Oliveira, of de evolution T., Peninsula. ps://doi.org/10.1111/evo.12177 influence the Italian L. changes the and of climatic Shirai, gies, case Pleistocene study A., Late The the Porto, deer? red Did the (2015). of G. size PalaeoecologyPalaeoclimatology, Sansalone, body in & trends L., O’Hara, evolutionary Maiorino, R., P. marsupial. L., Minchin, Pandolfi, D., large Sciences (2018). McGlinn, P. P., a Biological Solymos, Legendre, & R., in B: L., 2018 Kindt, G. M., Semelparity Simpson, Series Friendly, B., G., (2001). London. R. F. Blanchet, A. of J., Oksanen, Cockburn, Society & Royal J., the ps://doi.org/10.1098/rspb.2000.1369 A. of Bradley, dings M., marsupial Oakwood, carnivorous a of organization social Dasyuridae). quoll, and supialia: Spatial northern (2002). M. Australia. the Oakwood, northern of demography of and savanna for- ps://doi.org/10.1071/zo00028 Reproduction lowland bite (2000). the of M. correlates bats. Oakwood, Morphological (1997). phyllostomid (2009). of R. M. mandible ps://doi.org/10.25911/5d76398a4e2e7 L. and Monteiro, skull Oakwood, & the L., in A. diet https://doi.org/10.1111/j.1365-2435.2009.01549.x Peracchi, and investigate R., ce to M. tools complementary Nogueira, as methods morphometric Ecosystems ( Freshwater Geometric and dolphins (2019). common A. in Loy, variability & P., Nicolosi, threatened conservation. for a units’ of significant Nimmo, ‘evolutionarily contraction , & Defining M., range (1994). D. geographic Watson, C. G., Moritz, mediate E. rainfall Ritchie, Z., and C. ruggedness J. predator. Woinarski, Topographic marsupial E., (2019). L. Valentine, G. A., J. D. Dunlop, A., H. Moore, Feeding kangaroo, short-faced (2018). extinct dipro- giant S. extant among the Wroe, morphology . for craniofacial predictions & determines mechanics Dietary J., Biting herbivores: (2019). todont S. R. Wroe, & Flavel, R., (Ma- D. A., Mitchell, Pademelons Australian J. among Ledogar, scale subspecies G., Thylogale). the Sansalone, cropodidae: at morphology E., craniofacial determines influences Sherratt, foraging biomechanics R., of biomechanics D. The Mitchell, (2018). S. relatives. Wroe, their & and A., 285 kangaroos J. extant among Ledogar, in length E., shape face Sherratt, and R., size D. Mandible Mitchell, ecogeography. (2017). and A. taxonomy Loy, Research for & Evolutionary P., tool and A Ciucci, P., Mammalia): (Carnivora, Colangelo, Ursidae G., Guidarelli, C., geometry.Meloro, mandibular the by revealed Paleontology as Vertebrate carnivores of large Plio-Pleistocene Journal of habits Feeding (2011). C. Meloro, 9 Paleobiology 1) 373–375. (10), 18) 0885 https://doi.org/10.1098/rspb.2018.0845 20180845. (1881), . , 45 1,1711 https://doi.org/10.1017/pab.2018.46 167–181. (1), iest n Distributions and Diversity ora fZoology of Journal ora fMmainEvolution Mammalian of Journal , , h clg fteNrhr Quoll, Northern the of ecology The 55 n/a 4,2927 https://doi.org/10.1111/jzs.12171 269–287. (4), , epiu sp. Delphinus na.https://doi.org/10.1002/aqc.3042 (n/a). 440 , 31 1–1.https://doi.org/10.1016/j.palaeo.2015.08.038 110–115. , 2,4846 https://doi.org/10.1080/02724634.2011.550357 428–446. (2), , 257 ea:Cmuiyeooypcae akg eso 2.5-2. version package R package. ecology Community Vegan: 2,2728 https://doi.org/10.1017/S0952836902000833 237–248. (2), sn uemspecimens. museum using ) utainJunlo Zoology of Journal Australian , 25 rceig fteRylSceyB ilgclSciences Biological B: Society Royal the of Proceedings 12 1) 8813.https://doi.org/10.1111/ddi.12982 1818–1831. (12), https://doi.org/10.1007/s10914-018-9455-8 . ucinlEcology Functional Evolution ora fZooia Systematics Zoological of Journal qai osrain Marine Conservation: Aquatic rnsi clg Evolution & Ecology in Trends , ayrshluau.htt- hallucatus. Dasyurus 268 , ioteuu occidentalis Simosthenurus ayrshallucatus Dasyurus ayrshallucatus Dasyurus 67 , 48 16) 0–1.htt- 407–411. (1465), 1) 3532.htt- 3305–3322. (11), , 5,5959 htt- 519–539. (5), 23 Palaeogeography, 4,715–723. (4), Procee- (Mar- in , , Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. ieetpeoye rmtesm eoyei h aeevrnetb eeomna variation. of developmental Production by (2008). environment D. same C. Schubart, the & in J., Biology genotype O. Experimental same Schmitz, of the G., Dasyuridae). from Boogaart, (Marsupialia: M., phenotypes Thiemann, different Antechinus M., and Huber, G., Murexia Vogt, of revision Morphology-based Museum (2002). Memoirs-Queensland fossil S. the of Dyck, use and Van distribution. Australia, central past from its species trace new & a to J., of bandicoot, Cramb, record description pig-footed J., a the G. of with Price, Reassessment Chaeropodidae), D., sight: melemorphia, Stemmer, plain P., in Brewer, Hidden S., (2019). Brace, J. P., Louys, R. Miguez, Sim˜oes, F., J., B. K. Travouillon, combi- by raise in cetaceans to variation evidence for dental gical Investigating units (2016). Sinding, management J. J., Defining K. Travouillon, Nabe-Nielsen, (2015). M., J. tracking. Amundin, Teilmann, satellite C., & and J. https://doi.org/10.1016/j.gecco.2015.04.002 W., acoustics Koblitz, morphology, va- L. L., genetics, Size Andersen, Kyhn, ning monkeys: S., R., H. and Dietz, A., M. marsupials Galatius, neotropical S., in Sveegaard, modularity flexibility. Skull and (2010). constraint G. https://doi.org/10.1002/jez.b.21367 evolutionary Marroig, and & scar- riation T., in L. study the Shirai, A of (2018). Italy. implications C. Islands, conservation Corti, Pontine and & ps://doi.org/10.1093/biolinnean/bly091 western B¨ohme, W., differentiation the R., phenotypic Castiglia, of speciation, V., Incipient Avramo, P., let: impact Colangelo, foraging G., aquatic Senczuk, Does (2016). A. Herrel, & R., https://doi.org/10.1098/rspb.2016.1645 Godoy-Diana, snakes? in C., evolution A. shape Fabre, head R., Cornette, M., Segall, Eco- (1989). A. R. How, 7998.1989.tb02510.x & Australia. quoll, F., Western W. northern Kimberley, Humphreys, the J., teau, D. of Kitchener, physiology M., C. and (2019). Kemper, logy S. J., A. M. Bradley, Schlager, H., L. & debate. Schmitt, D., taxonomic Ian, a G., answer Jefferis, to S., Hungary Schlager, from remains moles humeral Electronica of Palaeontologia (2015). trajectories P. allometric Piras, and & metrics T., evolution. Kotsakis, G., mammalian Sansalone, early of relative in Conserved superimposition patterns (2008). optimal https://doi.org/10.1111/j.1525-142X.2008.00267.x S. ossification the Kuratani, 519–530. cranial & for O., of method Mock, timing V., Procrustes Weisbecker, the A., Goswami, of R., S´anchez-Villagra, Extensions M. (1990). D. Slice, & landmarks. monkeys: J., world F. old Rohlf, two in dimorphism sexual https://doi.org/10.1007/BF02547620 cranial 403. of ontogeny The cicularis (1991). J. M. Ravosa, Austria Vienna, Computing, (2019). Team. Core R Croihcne and (Cercopithecinae) ytmtcBiology Systematic :Alnug n niomn o ttsia optn.RFudto o Statistical for Foundation R computing. statistical for environment and language A R: .nst pallescens nasuta P. , 211 , 18 . 4,5053 https://doi.org/10.1242/jeb.008755 510–523. (4), rceig fteRylSceyB ilgclSciences Biological B: Society Royal the of Proceedings , aai larvatus Nasalis , 48 2,11.https://doi.org/10.26879/560 1–17. (2), 39 1,239–330. (1), 1,4–9 https://doi.org/10.2307/2992207 40–59. (1), Zootaxa ora fZoology of Journal ilgclJunlo h ina Society Linnean the of Journal Biological ayrshallucatus Dasyurus hms 93t pce rank. species to 1923 Thomas, , 4566 (Colobinae). ap fossilis Talpa 13 1,16.https://doi.org/10.11646/zootaxa.4566.1.1 1–69. (1), .Ep ol o.Dv Evol. Dev. Mol. B Zool. Exp. J. , 217 eaee nasuta Perameles lblEooyadConservation and Ecology Global akg ‘Morpho.’ Package nentoa ora fPrimatology of Journal International or 4,5958 https://doi.org/10.1111/j.1469- 539–558. (4), Mruila ayrde,a icelPla- Mitchell at Dasyuridae), (Marsupialia, ap europaea Talpa vlto Development & Evolution Zootaxa effo,10,wt morpholo- with 1804, Geoffroy, heou ecaudatus Chaeropus sn emti morpho- geometric Using ? , 4114 , , 283 125 , 314 4,351–392. (4), 13) 20161645. (1837), 1,5–0 htt- 50–60. (1), Podarcis , 8,663–683. (8), 3 aaafas- Macaca 839–850. , , , Journal 10 12 lizards (Pera- (4), (5), Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. Data may be preliminary. ero,T . afr,I . ree .(05.Semn h ie rgestwrsrsligthe resolving towards G., Progress E. northern tide: Ritchie, of the fauna J., mammal Stemming Smith, disappearing (2015). K., the Tuft, N. for E., responses Preece, Vanderduys, management & implementing K., Australia. J., and J. decline I. Webb, of Radford, Z., causes B., C. T. J. Woinarski, Reardon, Genetic R., (2016). M. G. Ziembicki, Amato, & S., ( Rabinowitz, jaguars management. Mesoamerican H., and of Quigley, structure I., population Dias-Freedman, and diversity A., Caragiulo, Carnivore C., of Wultsch, evolution the in constraint consistent remarkably and shape. Convergence skull (2007). N. Milne, & S., Wroe, characteristics. within male relationships and evidence Phylogenetic the molecular (2015). Mammalogy of on M. of based status Westerman, systematics conservation Quoll & Dasyuridae): C., the syuromorphia: Krajewski, and A., deposits P. Woolley, cave in specimens of of Zoology identity distribution present species the and recognised of Past Territory,currently (2013). understanding Northern M. Westerman, better Land, & a Arnhem A., Islands, Haslem, A., Company Distributional P. Woolley, (1999). English P. and Masters, & Wessel R., the Southgate, on K., Australia. Brennan, mammals A., of Fisher, C., patterning Palmer, Z., C. cause, Context, J. E. Australia: Woinarski, Ritchie, northern B., Gor- of Rankmore, fauna C., I., mammal S. Radford, R. C., disappearing Palmer, Firth, The response. L., A., (2011). and N. Fisher, M. McKenzie, Ziembicki, A., N., & A. C. S., Burbidge, Ward, fau- Johnson, J., G., D., B. continental A. Traill, Griffiths, A., a J., J. I. of Fitzsimons, don, S., unraveling Legge, Un- Z., Ongoing C. settlement. (2015). J. recovery: Woinarski, European L. since P. tiger mammals Harrison, Australian Sciences Planning & N., of of A., Academy extinction (2015). Balkenhol, A. and C. Burbidge, Decline L., na: Z., Orlando, A. C. K., J. Kitchener, Woinarski, A. conservation. & Scharf, effective J., J., for Fickel, ps://doi.org/10.1126/sciadv.1400175 variation Niedballa, S., intraspecific P., Kramer-Schadt, derstanding Christiansen, do- H., A., system Hofer, Courtiol, masticatory Col- the wombats. A., E., of marsupial C. Wilting, variation Terhune, herbivory-adapted C., Individual the A. (2019). in Sharp, O. shape M., https://doi.org/10.1186/s12983-019-0338-5 Panagiotopoulou, skull H. & Abraha, 3D E., S., minates Sherratt, Johnston, C., in S., Speck, heterochrony lins, T., Ossification Guillerme, dichotomy. (2008). V., Weisbecker, marsupial-placental R. the M. and Sanchez-Villagra, skeleton & https://doi.org/10.1111/j.1558-5646.2008.00424.x (2019). postcranial S., A. Wroe, therian Goswami, A., the & Goswami, A., V., Herrel, integration. M¨uller,Weisbecker, cranial J., A., of Sciences J. pattern of Academy conserved Maisano, National from N., the squamates R. in Felice, diversification C., MorphoEvoDe-Ecomorphological A. skull. the Fabre, of A., Integrating form Watanabe, the to muscle morphology! temporal the of live relation phylogenomics. , The (1947). deadLong L. and S. Washburn, EvoDevo is molecular Morphology ps://doi.org/10.3389/fevo.2015.00054 into (2015). vo A. Wanninger, 99 3,2928 https://doi.org/10.1002/ar.1090990303 239–248. (3), , 61 Therya Zoology of Journal Australian Evolution 4,2120 https://doi.org/10.1071/ZO13034 281–290. (4), osrainLetters Conservation , 96 LSONE PLOS , 6 1,3–6 https://doi.org/10.1093/jmammal/gyu028 37–46. (1), 1,169–225. (1), , , 112 61 5,15–20 https://doi.org/10.1111/j.1558-5646.2007.00101.x 1251–1260. (5), 1) 5144.https://doi.org/10.1073/pnas.1417301112 4531–4540. (15), .blythi D. , 11 , , 1) 0637 https://doi.org/10.1371/journal.pone.0162377 e0162377. (10), 116 4 and 3,1221 https://doi.org/10.1111/j.1755-263X.2011.00164.x 192–201. (3), 2) 48–49.https://doi.org/10.1073/pnas.1820967116 14688–14697. (29), , 47 .cristicauda D. 1,8–1.https://doi.org/10.1071/ZO99004 87–111. (1), 14 rnir nEooyadEvolution and Ecology in Frontiers Mruila Dasyuridae). (Marsupialia: ateaOnca Panthera cec Advances Science rnir nZoology in Frontiers Evolution :Ipiain o conservation for Implications ): rceig fteNational the of Proceedings , h ntmclRecord Anatomical The 1 utainJunlof Journal Australian , Dasycercus 5,e407.htt- e1400175. (5), 62 8,2027–2041. (8), Dasyurus rceig of Proceedings , , 16 3 Toward : Journal 1,41. (1), htt- . (Da- Posted on Authorea 7 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158359716.69973386 | This a preprint and has not been peer reviewed. 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