Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. hnnah Munyai Thinandavha encroachment bush to responses confounds assemblage custodiens) ( ant Pugnacious A ihes(oh dad,Bacehr,Wle,&Hfr 05.Hrn ta.(09 n lu tal. et Blaum at richness and and (2019) abundance in butterfly al. increasing affecting families et in some not Hering vary with but cover beetles, 2015). cover, on shrub effects Hofer, shrub of mixed & intermediate change similar gradients Walter, habitat at reported However, Blanckenhorn, (2009) local peaking by Edwards, richness affected 2009). (Koch, worst grasshopper Jeltsch, richness the with & be impact, will Schwager, ability their Rossmanith, dispersal low Seymour, with (Blaum, those conservation and for Eswatini implications Wingless of real diversity. has Kingdom maintain This the to plant in 2012). aim growth communities woody Monadjem, that herbaceous & bird by (Sirami managers while of affected structure 2008) richness in strongly Morgan, species shifts & significant is Australia, showed (Price southern reduced and forage Park, Overall, significantly functioning production was National reduce forms 2017). ecological Grampians also in al., both encroachment Bond, et jeopardizing e.g. bush encroachment, (Archer & areas, and protected ectoparasites Gray, habitat, and increase (Parr, the lands and Africa biodiversity of grazing quantity communal of threatens In and rest encroachment quality services. the plant the across woody year per reducing that 1.275% per 2012), predict to 0.25% -0.131 and studies between 2014), Broad-scale varied 2017). Hoffman, have Durigan, & Over rates & Puttick, Murphy, encroachment 2020). (O’Connor, Lehmann, 2017) Stevens, (Stevens, Africa & al., South Lehmann, et in grasslands, Bjorkman, (Archer and a year Myers-Smith, savannas centuries pose into (Criado, two encroachment 2018) plant particular past al., woody open in the of a et change largely that (Osborne understory, climate is a threat activities to considered pervasive grassy anthropogenic linked One therefore, continuous various and, biome. However, by this 2019), to 2020). characterised Bond, threat Bond, 2017; is & al., and (Pausas et ecosystem Africa, (Archer overstory southern woody dominates discontinuous biome savanna species. The this of dominance indiscriminate behavioural they disturbance, and to where numerical Introduction positively the soils, responds with clayey also associated with custodiens effects habitats A. on open As knock activity. in to ant abundant lead all locally could of were clearing were 90% characteristics bush that than site-specific more species These for a in accounted important. variation custodiens, often of more Anoplolepis amounts were species significant to seasonality Although explained linked and Thickening thickening. characteristics co-occurring largely lower. woody two site-specific significantly in extensive but was experiencing evenness epigeic composition, thickening, biome habitats, of ant woody savanna open response to African in the response an higher study in vegetation of was to ants horizontal richness traps open) for pitfall and and true used vertical (closed particularly We increased habitats is diversity. of ant This dominant on effects communities. impacts The negative in biodiversity. predicted shifts with in marked variation in of result determinant can key structure a is structure Habitat Abstract 2020 25, November 2 1 nvriyo wZl-aa olg fArclueEgneigadScience and Engineering Venda Agriculture of of University College KwaZulu-Natal of University 1 tfnFoord Stefan , 2 o Slotow Rob , 1 1 n oahmaq Mkhize Nomathamsanqa and , 1 Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. fe olcigdt,smlswr ahdi h aoaoy n h pcmn one,sprtd and separated, m 10 counted, least specimens at the were and traps laboratory, pitfall the the and in grid washed in sampling were traps 2 samples pitfall X data, ten 5 days. collecting were a five After a There in for as and out open 2012). glycol laid season) left Foord, propylene were were (wet-hot & of traps and 2017 (Munyai solution apart, pitfall January ants 50% The in repels contained conducted plot. described (62mm) nor each was as traps attracts site Pitfall neither method each that trapping season). at preservative (dry-hot pitfall Sampling 2017 standardized October a (2015). early Foord following and sampled Munyai were in ants worker a dwelling with a Ground habitat-type with (2012), habitat-type al. and savanna, et sampling habitat. Canopy open Parr Ant closed grids. as in 48 as middle treated were described treated the were there were As in total 3% 55% were In type. [?] plots edge. [?] habitat of the habitat the cover from least closed away canopy categorize The m at mean site. to 30 were were per used replicates plots site was open eight cover the each of (Mag), while total within habitat, Magangeni a this Replicates for (Isi), of respectively), Isivivane (Zin). open (Gon), Zincakeni and Gontshi closed and study: (Mem), this gate > for Memorial selected (Man), were Mansinya sites sampling Six-paired wildlife a woodland selection is savanna Site and 2013). open Wildlife, Bond, to (KZN) KwaZulu-Natal & Reserve) Ezemvelo habitats (Gray Game by 2010). of managed area (Hluhluwe Midgley, majority is conservation side & park The northern thicket. The (Boundja Reserve). shrub the October-March Game with on km between (iMfolozi interspersed forest savanna summer (900 woodland semi-deciduous in with Park from covered usually vary is Park mm, park Hluhluwe-iMfolozi 985 the (28.2198 of to of Most Africa mm section 700 South Hluhluwe from KwaZulu-Natal, ranging the northern in Zululand, conducted in was study The soil seasonality, how area (1) Study experiencing differences. ask biome these and savanna with African whole, Here, associated an assemblages. a species Methods in ant identify as diversity in and ant community (2) abundance taxa affect thickening, the dominant species to woody of on interact ant extensive result complexity focusing model the habitat by to largely structural study extensions and is multivariate roles their Although their functional complement Ant taxa. and we 2012). rare in models 2012). Wang, al., these & linear (80% Wright, et on (Warton, generalized (Warton predatory composition focus community and use reflect largely location mainly not we confounding were to results to shown Here, more Their taxa leads been that has used habitats). restricted and approach observed effects open distance-based these dispersion but the in differences, that Park, 7% compositional Hluhluwe-Imfolozi and the analyzed in they to habitats between habitats compared closed differences closed forest real to varies the and no restricted and found open (2012) (Del investigated were in al. services been et species activity Parr richness ecosystem rarely and However, increased and 2005). and richness has 2009) (Bestelmeyer, 2018) ant al., encroachment USA et western Landis, (Blaum to south Kalahari & the southern response in the (Wills Their in abundance functions 2012). increased from ecological Pelini, geographically of & myriad Ribbons, a Toro, (H arthropods perform terrestrial abundant and most 2009). the 1990), among al., example are Formicidae), et for (Blaum (: arthropods, Ants decomposition Morente-Lopez, by dung Garcia, provided adverse Lara-Romero, and 2011) have 2017; 2016), al., will Galen, Iriondo, et encroachment & & (Eldridge plant Moffett, services woody Miller-Struttmann, ecosystem abundance that (Kettenbach, and their is pollination affect ecological consensus not the general did on However, Australian but effects in 2016). composition, species community Eldridge, shrub & e.g. of (Kwok differently, drivers respond important also were might woodlands metrics Different levels. encroachment higher 0maatwieteata ie eea es maat ahst a orpie lt oei the in (one plots paired four had site Each apart. km 2 least at were sites actual the while apart 300m 2 ° ,31.9519 S, ° ) h akrcie rainfall receives park The E). ca. encnp cover canopy mean ldbe Wilson, & ¨ olldobler 2 ,situated ), ca. Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. adm R random, epne fatasmlgst rdet nsi n aia tutr eemdle sn Threshold using modelled were structure habitat and soil in gradients to samples visualization, assemblages aid ant To of 2016). 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The to complex axes soil %. used more two and 27.36 were and second first litter, components the with The leaf principle and associated cover, These 41.92, positively canopy S1). axis, was increased (Figure first PC1 to variation, soils %. related the clay 14.57 positively of second with explained % was the soils 69.28 associated for and explained negatively PCA %, structure the 51.98 and habitat of principle axis soil axes a first two sandy the first doing variation, The more by the respectively. for variables of accounted of % was sets 66.55 two structure) the habitat of and analysis and (soil component Research moisture). variables % predictor the manganese, and between to calcium, Collinearity silt sent % potassium, were sand, (phosphorus, % samples clay, content a Analysis soil (% chemical using Statistical texture 48 clay), collected and of (% % copper), were total analysis and and samples composition magnesium A nitrogen subsoil for zinc, measure Pietermaritzburg, % 10 a replicate. 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Data may be preliminary. epne ohbttsrcuecagswr o rda Fgr 4.Ti seiecdb h eaieylar- relatively the by evidenced is This characteristics, S4). soil (Figure in gradual gradients spe- to not more responded were ants significantly changes although had structure and, habitat habitat habitats; Open to closed composition. than responses and even richness less ant were but (open on cies, type impacts habitat important structure, had assemblage closed) ant dominated vs seasonality and characteristics site-specific Although Discussion types. habitat open species, and two ( closed while 18 the types, both habitat in open with sampled associated mostly and sites, these, all of Three across distributed were species Five of while eight habitats, activities closed while The 3). and structure, habitats. soror open (Figure habitat closed both site and in in study soil um open gradients the sandy across the in more soils in gradient of variation clay increased sand-clay any marginal the the of with preferred with amount associated tably largest associated principle were soil third were first the taxa taxa the explained indicator no although S2), Similarly, variables, Figure three 2). see significant previous (Table explained soil, the variables sandy by soil (increasing explained conside- and when structural that component the particularly beyond of variation composition, None of species 2). amounts in (Figure variation explained dis- most variation the conditional the ring explaining by ant in exemplified in seasons was followed variation Habitat assemblages of on amount sites largest of of impact the comprised and This 1). explained assemblages (Table 2). ant study characteristics Figure site-specific tinct this 2, while i.e. (Table in observed, factor, structure variation random measured assemblage the the not of to 22% characteristics linked also around was was Site-specific variation explained Soil-pc1 this models modelled. of three was half 0.026 dominance All model, least = Simpsons model. second at (estimate when second-best the pronounced soil the in more sandy included in even 0.014 to was included became = Soil-pc1 response habitats richness, positive (estimate open to a soils significantlyof Similar for was sandy habitats. there evidence soil more that open weaker included evidence in the with strong model richness in with second-best no.1, diversity in The Hill signifi- Shannon increase considering 1). less being when an (Table reversed richness season for was with wet trend evidence this the (closed/open), However, weak during habitat very and and with habitat season open PC1, included the richness in species higher cantly for model best The genera specious most The ants. subfamilies. genera). five six and and species, genera, species Poner- 25 by (15 followed in subfamily, siota and species abundant and genera), 74 diverse eight comprising most were and the collected, species was were (13 genera) specimens eight inae and ant species 008 (47 61 as Myrmicinae well 2010). of King, as total & bootstrapping, A (Baker through points classification change same community the and given taxa Results is individual taxon of a location those the that from around Mendez- times distinguished uncertainty Pardo, of are Costas, proportion 2010; (z+) the King, responses and & Increasing (Baker uses gradient ( 2018). method a decreasing Rodriguez, This along & response 2010). of Martinez-Madrid, direction King, Fernandez, the & and (Baker points, package change “TITAN2” the from standardized (TITAN) Analysis Taxa Indicator UKZN Lepisiota squaminode Tetramorium and , and npoei custodiens Anoplolepis oooimjunodi Monomorium 2( 22 Pheidole z- z UKZN taseiccag on Bkr&Kn,21) IA loetmtsidctrreliability indicator estimates also TITAN 2010). King, & (Baker point change specific a at ) soe bandfo niao pce nlss(niao au)t eettespecies-specific the detect to Value) (Indicator analysis species indicator from obtained -scores gabonense Pheidole p)wr soitdwt lsdhbtt(iueS3). (Figure habitat closed with associated were gp.) 2 species), (22 a v pce ah he ureso l h n niiul agtblne oone to belonged caught individuals ant the all of quarters Three each. species five had 2( 02 UKZN capensis p)hdterhgetatvte nca soil. clay in activities highest their had gp.) hsseis n hs ntegenus the in those and species, This . ciiispae nlaysoils. loamy in peaked activities 1( 01 Monomorium p) soitdwt h ad ol fMniy Fgr iueS1). Figure & 2 (Figure Mansinya of soils sandy the with associated gp.), megacephala dnoahstroglodytes Odontomachus and gp.), Pheidole 4 eooeacaffraria Mesoponera hioecasnd ruspolii crassinoda Pheidole ahhdsxseis while species, six had each , Tetramorium Pheidole npoei custodiens Anoplolepis ymcranatalensis Myrmicaria and ± cone o 0 falthe all of 90% for accounted , Monomorium UKZN .5.Tengtv impact negative The 0.05). dnoahstroglodytes Odontomachus and Tetramorium 4( 14 Camponotus .custodiens A. simillimum and UKZN , Bothroponera Tetramori- ± UKZN 3were 03 0.015). , Lepi- gp.), no- - Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. loaetdoealatdvriypten ntersuy ons or,adVle 21)as observed also (2014) Villet and have Moore, could Bownes, turn study. in which their ants, in of dominant patterns number the soils diversity large sandy of very ant that abundance suggested overall the study favoured affected latter have The also soils. could sandy Valiente-Banuet, complexity in Rios-Casanova, habitat abundance Mexico, ant and central Tehuac´an high of the reported Valley (2006) of Rico-Gray fan and alluvial an Along of soils. activities loamy contrast, abundance) ( In total species variation 3). the other of (Figure of driver soils % the (72 clay India. hyperabundant as with Eastern the (2015) of of Basu habitats activities and forested The the Chatterjee, in Parui, composition in vegetation assemblage reported in ant were changes that litter, in overall complexity, noted properties than vegetational is (2018) chemical namely, surface Eldridge measure characteristics, soil soil and the site-specific not and Gibb, on of did Radnan, alteration influence small-scale general, we The to In composition. that study. responsive community variables this more these of be or Whether to scope autocorrelation tend 1). the ants (Table beyond spatial effects and predictors factors, marginal say influential historical and to most difficult conditional the to were both related own by their are suggested on differences as specific sites structure, Similarly, with assemblage soil S2). sandy ant (Figure on of site is site-specific this site of Mansinya with importance The associated the characteristics. on species soil highlighting to therefore related and that assemblage particularly ant characteristics, distinct a had Mansinya, site, 2015), One Bond, & is Midgley, Park Staver, habitat. Hluhluwe–iMfolozi also (Charles-Dominique, open the could type crucial dominate habitat vegetation a Savannas open savanna as the open covering open in, Currently, in by there effect. species references dominated of species-area emphasises and number largely again, the (2019), larger once by Andersen this This, only explained However, in habitat. were composition. be closed detailed species assemblage to the as which ant restricted of openness, Feldhaar, rugosity, of % % habitat & driver 24 8 surface of total, the Rothmaier, ground (Campos, role In to Teuber, complex. important of compared composition Blaha, more habitat, the effect and structural open (Greve, largely the the richness speed is in of habitat assemblage running recorded closed because ant ant study, largely this example the In is for 2019). on This activities, effect 2006). foraging clear Ribeiro, affects variation a overall & and have cover Martins, plant habitat intensity, must Soares, sunlight of that of structure expected amount effects however, the habitat is, the example It for in different. reported site, significantly a (2019) were in habitats al. characteristics two natural woody the et or in assemblages Guilherme stems behaviour. ant grass surprisingly, basin, and Not by strategies Amazon impeded foraging the structure usually species habitat are In ant that they 2019). given on as surprising, al., complexity ants not of et this is movements (Nooten This of the . determine twigs assemblages 2012) to ant Foord, thought the & amount been (Munyai significant in had assemblages a variation ant explained types conditional in warmer vegetation variation of mountains, richnessof and Soutpansberg amount western species open largest the ant across forest. more third Similarly, study. secondary of the reported or explained response (2019) primary habitat negative compared al. Overall, a the et than (2004), reported Hethcoat ants Hochuli (2019) Similarly, more and complexity. al. supporting Lassau habitat pasturelands contrast, et of In with Ahuatzin gardens Cerrado. species diversity urban and Brazilian fewer and altered (2016) the and reported highly Foster of also flora in habitats (2012) and was Vasconcelos both complex (2019) Yusah aite and structurally of Cook Pacheco study least and driver Australia. their the Facey, south-east Notably important in Rowe, of habitat. Schultheiss, an Region open Nooten, Sydney is in study, Greater cover species our vegetation ant to fewer of contrast reported amount in However of the communities. presence that fauna the reasoned by (2019) explained be Andersen partly thickening can to response Responses traglodytes threshold diversity. Odontomachus This ant explaining abrupt. dominant in numerical be closed) the to versus predicted (open therefore, classification are, categorial of role ger > 5 ftelnsae hsi tefwudepantelre ubro pce bevdi the in observed species of number larger the explain would itself in This landscape. the of 75% ymcranatalensis Myrmicaria npoei custodiens Anoplolepis .custodiens A. hc saseilsda ela grsiepeao nsnysoils. sandy in predator aggressive as well as specialised a is which , ohooeasoror Bothroponera noe aiaso oeststa eemil ncae ol and soils clayey on mainly were that sites some of habitats open in ni irsocad fteEsenCp,SuhArc while Africa South Cape, Eastern the of orchards citrus in in dnoahstroglodytes Odontomachus 5 and oooimjunodi Monomorium npoei custodiens Anoplolepis ekdi ad ol.Three soils. sandy in peaked ,wr oeatv in active more were ), nrae nsite in increased , A. Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. nesn .N 21) epne fatcmuiist itrac:Fv rnilsfrunderstanding for principles Five disturbance: group. to faunal dominant communities globally ant a doi:10.1111/1365-2656.12907 within of of remnants dynamics Responses C., rainforest disturbance (2019). M. the primary N. Ribeiro, in M., A. diversity R. ant Andersen, Feitosa, litter E., leaf J. drives Gonzalez, cover 019-01712-z Forest Valenzuela (2019). A., landscapes. W. A. tropical human-modified Dattilo, Jaimes, . J., . E. vineyards. Corro, in A., forage that D. Formicidae) Ahuatzin, (Hymenoptera: Africa. ants South of Province, survey African A Cape the theWestern (2000). of J. in M. potential Samways, invasion & P., the Addison, demonstrate habitats Surrogate (2006). ant. M. pugnacious Samways, & P., Addison, authors. References the the among collected interest NNM of and article.. conflict TCM the No paper. revised SHF Statement the and Interest wrote RHS of and NNM, Conflict data TCM, the ants. analysed the study, identified TCM the data, designed SHF and TCM Khumalo. Eric and Dlamini Falakhe assistants contributions guards, Field Gun- Author Park. game Xaba, Hluhluwe-iMfolozi Phethele Hluhluwe-iMfolozi at Mutavhatsindi, the research Joshua Ezemve- and this Kunene, the Makhuvha Caroline conduct and do Xolo, to 114416) Sbongiseni No. permit (Grant Thabethe, a author Nokubonga us first from granting the paper. for to this wildlife funding of KZN for Funding lo publication Research the National regarding the interest thank We of conflicts on declare authors The predicting around better note interest a cautionary of provide patterns. a should Conflict global this tool, broad-scale consider management on we a based as though, impacts mammals Generally local browsing impacts. or possible fire of using understanding shrubs, and trees of of removal response and the 2000), 2008). (Mitchell, Exploring Parr, species specific. ( ant savanna other African custodiens displacing South A. 1990), the al., in et since structure Prins Particularly assemblage 2018; services. ant this al., ecosystem control facilitate et that also could (Mauda fact but vertebrates sites diversity, The small of ant indiscriminately. clearing on done 2006), only observation), Samways, be not Personal custodiens & not impacts Munyai (Addison should negative (TC habitat with shrubs particular open invasion and dominate in completely trees soils can shallow of species removal open manual in that supercolonies 1992). suggests of (Dean, formation systems the agricultural the with in 2000). pests Samways, custodiens serious & A. Addison the other 1990), 2006; of of Samways, one Prins, variety & as & a (Addison regarded on Robertson, landscapes hence, prey (Prins, agriculture to in livestock known pest small well major 2008; custodiens also is a Parr, and it also 2018; 2015), region, is Foord, the Pruitt, it & to & Munyai, native Wright, Seymour, is (Keiser, Joseph, species invertebrates (Mauda, this Although studies of 1983). African Samways, impact southern and several behaviour, in dominant mented abundance, high The custodiens sas elkonfrtnighnye rdcn etisc pce,adbignca collectors, nectar being and species, pest producing honeydew tending for known well also is a enosre ons noe n elisltdsiso h ao Da,1992). (Dean, Karoo the of soils insulated well and open in nest to observed been has saplgnu Smas 90,gnrls rdtro te netbae Da,19) and 1992), (Dean, invertebrates other of predator generalist 1990), (Samways, polygynous a is spriual ciei lysis n laigo od pce ntepr edt econtext be to need park the in species woody of clearing and soils, clay in active particularly is epn oiieyt itrac Adsn&Smas 00 ad ta. 08.Coupled 2018). al., et Mauda 2000; Samways, & (Addison disturbance to positively respond idvriyadCnevto,15 Conservation, and Biodiversity .custodiens A. idvriyadCnevto,28 Conservation, and Biodiversity fia noooy 8 Entomology, African ovroswoyseiscnrlmaue,b tmanual it be measures, control species woody various to 1,4148 doi:10.1007/s10531-005-5399-4 411-428. (1), 6 .custodiens A. ora fAia clg,88 Ecology, Animal of Journal 251–260. , 5,19-17 doi:10.1007/s10531- 1091-1107. (5), natdvriyhv endocu- been have diversity ant on Acknowledgments 3,350-362. (3), A. A. Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. lrde .J,Bwe,M . asr,F . oe,E,Ryod,J . htod .G (2011). G. W. Whitford, (2016). Press. & B. California F., of Bolton, University J. & synthesis. L., Reynolds, global B. a E., towards Fisher, Roger, functioning: T., and structure F. ecosystem 14 Maestre, review on Letters, encroachment a A., shrub revisited: M. world of Bowker, the Impacts J., run that D. things Eldridge, little The Formicidae). Southern (2012). (Hymenoptera: the L. disservices S. in and 133-146. Pelini, services species & ecosystem ant R., ant-mediated R. some of Ribbons, of I., patterns Toro, Woody activity Del (2020). determining N. Temperatures Africa. Stevens, (1992). South & J. Karoo, R., R. E. W. C. biomes. Dean, savanna Lehmann, and D., tundra across A. change Bjorkman, 29 climate Biogeography, H., under intensifies I. encroachment Myers-Smith, plant G., of M. Sensitivity Criado, (2018). Spain. P. Northern differentia- in Rodriguez, areas & Functional mining in M., gradients (2015). 93 Martinez-Madrid, metal to J. L., taxa Mendez-Fernandez, indicator W. macroinvertebrate I., mosaic. Pardo, Bond, savanna/forest N., & Costas, African ant F., an on G. structure in Midgley, habitat doi:10.1016/j.sajb.2015.05.005 biomes of C., of Effect A. (2006). tion Staver, P. T., S. species. Ribeiro, Charles-Dominique, tree & pioneer P., two R. to webs. associated Martins, hemipteran-tending foraging 737. Formicidae) P., recruiting of : J. partitioning enemies: (Hymenoptera Soares, assemblages enemy’s spatial I., by My R. orchards (2014). Campos, citrus H. in M. control Hluhluwe- Villet, 22 biological the & Entomology, for in D., ants plants S. generalist woody Moore, on A., impact Bownes, elephant of Patterns (2010). J. Ants,2028.2009.01104.x Africa. (2006). South J. L. Kwazulu-Natal, Midgley, S. park, Chown, & Imfolozi & Region. P., P., Floristic H. R. Davids, Cape Boundja, A., northern Niekerk, the Van in G., change H. and Robertson, altitude A., M. McGeoch, A., Botes, indicators. suitable Press. of Univ (2019). identification J. rangelands: savanna W. in Bond, diversity cover arthropod shrub 18 in of Changes Conservation, gradient (2009). and diversity driven F. Biodiversity Jeltsch, use & ant land M., Schwager, a of E., along Rossmanith, Enhancement C., Seymour, N., biodiversity? Blaum, diminish USA. desertification south-western in Does 9516.2005.00122.x invasion ecological (2005). and shrub biodiversity T. by interpreting and B. detecting for Bestelmeyer, method new Woo- A (2017). (2010). R. S. R. S. thresholds. community King, Woods, & (Ed.), & E., Briske J., M. D. R. Baker, D. Steidl, S., In Schwinning, Challenges Consequences. I., and and K. Management Causes Predick, Encroachment: M., Plant E. dy Andersen, R., S. Archer, 0-1.doi:10.1016/j.ecolind.2018.04.059 207-218. , 7,7972 doi:10.1111/j.1461-0248.2011.01630.x 709-722. (7), 3,5959 doi:10.4001/003.022.0303 519-529. (3), 5,9593 doi:10.1111/geb.13072 925-943. (5), pnEoytm:EooyadEouinbyn h oetEdge Forest the beyond Evolution and Ecology Ecosystems: Open ora fteEtmlgclSceyo otenArc,55 Africa, Southern of Society Entomological the of Journal ehd nEooyadEouin 1 Evolution, and Ecology in Methods p.25-84). (pp. 5,18-19 doi:10.1007/s10531-008-9498-x 1187-1199. (5), nso fiaadMdgsa:agiet h genera the to guide a Madagascar: and Africa of Ants iest n itiuin,11 Distributions, and Diversity fia ora fEooy 48 Ecology, of Journal African 7 ot fia ora fBtn,101 Botany, of Journal African South 1,2-7 doi:10.1111/j.2041-210X.2009.00007.x 25-37. (1), ora fBoegah,33 Biogeography, of Journal 1,2624 doi:10.1111/j.1365- 206-214. (1), 1,4-5 doi:10.1111/j.1366- 45-55. (1), agln ytm:Processes, Systems: Rangeland ymclgclNw,17 News, Myrmecological oibooy 47 Sociobiology, 1,149-156. (1), clgclIndicators, Ecological e ok Oxford York: New . lblEooyand Ecology Global 71-90. , ntdStates: United . 3,721- (3), 82-90. , African Ecology , Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. uyi .C,&Fod .H 21) nso onan pta,evrnetladhbttassociati- habitat and environmental endemism. spatial, of mountain: centre a a on in Ants transect (2012). altitudinal H. (Eds.), an S. Panizzi along Foord, R. ons & A. & C., Schaefer T. landuse W. Munyai, in C. Changes In (2018). (Coreidae). importance bugs H. economic Leaf-footed S. of (2000). Foord, L. & P. C., savannas.Mitchell, T. African in Munyai, groups L., 27 functional C. dominant Conservation, and and Seymour, composition S., assemblage diversity, G. ant alter Joseph, assemblages. ant V., on E. complexity Mauda, habitat of Effects (2004). F. doi:10.1111/j.0906-7590.2004.03675.x D. Hochuli, 157-164. & A., S. shrub on Lassau, of density effects plant interactions. indirect and visitor fine-scale Direct plant-flower and (2016). by doi:10.1111/1365-2435.12637 M. mediated 1521-1530. species J. grasslands shrub Iriondo, alpine & of on J., encroachment influence Morente-Lopez, C., The Garcia, (2016). C., Lara-Romero, J. shrubland. D. semiarid Eldridge, a & in C., arthropods B. pastures. subalpine A. affects encroachment Swiss Kwok, Shrub two (2015). alpine on G. grasshoppers the Hofer, 47 and & Research, using butterflies, T., Alpine Walter, study plants, U., of case W. diversity Blanckenhorn, A the J., P. wildflowers: Edwards, under alpine B., encroachment Koch, for shrub How services (2017). pollination viscosum. C. Polemonium Galen, threaten skypilot, & could Z., change Moffett, N., climate can Miller-Struttmann, colonies A., spider J. Social size. Kettenbach, group societies: increased arthropod and Warring apparency (2015). reduced N. via J. 119 ants Pruitt, predatory & by M., annihilation delay C. Wright, N., C. R. Keiser, diversity. in and data Entropy abundance (2006). multivariate L. of Jost, analysis 7 regression Evolution, and and Ecology ordination in boral–Bayesian Methods (2016). K. F. Hui, dad,D .(09.Teipc fscnayfrs eeeaino ruddeln n communities ant ground-dwelling A., on F. regeneration Edwards, forest H P., secondary C. of F. Achiardi, impact Andes. M., The Tropical C. (2019). the P. Ortiz-Sepulveda, in D. F., Edwards, F. a . Castiblanco, . J., along en- . B. niches Shrub King, G., (2019). species M. N. Hethcoat, beetle savanna. Blaum, ground-dwelling Namibian & B. semi-arid from M., a Insights F. doi:10.1002/ldr.3197 Schoenen, in degradation: Baccaro, gradient A., land cover & Marquart, always shrub K., C., not as- Geissler, A. is M., ant croachment Chagas, Hauptfleisch, ground-dwelling das R., of P., Hering, Basin. composition Pequeno, Amazon trait E., the functional Franklin, across doi:10.1016/j.actao.2019.05.004 P., predict test A L. complexity surface semblages? ground environmental J. of Can Souza, effect (2019). R., The (2019). dependent. D. and size H. Guilherme, than Feldhaar, experience rather & visitor species-specific M., the is Rothmaier, doi:10.1007/s00040-019-00694-z speed impact J., 364. running encroachment Teuber, ant plant S., on Blaha, rugosity woody E., Will M. (2013). Greve, J. W. areas? conservation Bond, of & economy F., E. Gray, ldbe,B,&Wlo,E .(1990). O. E. Wilson, & B., ¨ olldobler, 42.doi:10.1016/j.beproc.2015.07.005 14-21. , 2,3537 doi:10.1657/aaar0013-093 345-357. (2), 4,9795 doi:10.1007/s10531-017-1474-x 947-965. (4), eooi,191 Oecologia, lrd:CCPress. CRC Florida: . ode 55 Koedoe, clg n vlto,7 Evolution, and Ecology agln ora,38 Journal, Rangeland 6,744-750. (6), 2,4542 doi:10.1007/s00442-019-04497-8 475-482. (2), io,113 Oikos, h ants The 1,9 doi:10.4102/koedoe.v55i1.1106 9. (1), caOclgc-nentoa ora fEooy 99 Ecology, of Journal Oecologica-International Acta ascuet:HradUiest Press. University Harvard Massachusetts: . 2,363-375. (2), 8 adDgaain&Dvlpet 30 Development, & Degradation Land 1) 9367.doi:10.1002/ece3.3272 6963-6971. (17), 4,3139 doi:10.1071/rj15019 381-389. (4), ora fIsc osrain 16 Conservation, Insect of Journal netsScax 66 Sociaux, Insectes ucinlEooy 30 Ecology, Functional eaiua Processes, Behavioural rtcAtrtcand Antarctic Arctic cgah,27 Ecography, Biodiversity Heteroptera 1,14-24. (1), 677-695. , 3,355- (3), 27. , (2), (9), Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. iai . oajm .(02.Cagsi idcmuiisi wzln aansbten1998 between savannas Swaziland in communities bird in encroachment. Changes shrub to 4642.2011.00810.x (2012). owing A. 2008 Monadjem, and & habitats abundance C., of for Sirami, management series and assemblages a ant epigaeic in scarcity. variability: Formicidae) temporal and Species (Hymenoptera: (1990). ants M. Samways, of structure citrus. Community with with associated (1983). relationship Mexico. Valley, its M. Tehuacan and Samways, the food diversity of on fan Ant 29 effect alluvial Ecology, larger an of (2006). in a Journal V. factors Rico-Gray, has soil & complexity and vegetation A., surface Valiente-Banuet, Soil shrubland. L., to Rios-Casanova, (2018). grassland J. from D. change southern Eldridge, a of doi:10.1111/een.12510 than & (Eds.), areas ants H., agricultural Cedeno by and Gibb, A. exploitation urban N., & in G. Jaffe, ants Radnan, K. Pest herb-rich Meer, (1990). of der A. richness Press. Van Westview Prins, species R. Boulder: & reduces G., In encroachment H. plant Africa. Robertson, Woody J., A. (2008). Prins, W. J. Australia. southern Morgan, in Ecosystems. & assem- Terrestrial woodlands N., in species States J. Biome ant Price, Alternative shaping (2020). India. characteristics J. W. Eastern Habitat 25 Bond, Science, & in G., (2015). J. forest Pausas, P. deciduous Basu, mixed of & consequences a doi:10.1017/s0266467415000036 S., functional in and Chatterjee, biodiversity blages K., Cascading A. (2012). Parui, J. savanna. W. African switch. Bond, South biome & a 4642.2012.00882.x change-induced F., in richness E. global Gray, species a L., assemblage C. control Parr, can ants 77 Ecology, Dominant Animal of (2008). Journal L. heterogeneous naturally C. a Parr, in diversity ant enhances diversity Habitat landscape. (2012). Brazilian H. Vasconcelos, (2018). & R. R., E. Pacheco, C. Lehmann, traits. functional & plant G., by Midgley, mediated J., are doi:10.1111/nph.15236 W. savannas changes Bond, African Africa: complexity in N., southern impacts Stevens, Habitat Human in T., encroachment Charles-Dominique, P., Bush (2019). C. (2014). Osborne, Formici- M. T. M. J. (Hymenoptera: Hoffman, & spaces Cook, R., causes. green & J. and Puttick, urban L., G., in T. S. assemblages O’Connor, Facey, ant C., of R. diversity and Rowe, dae). traits P., functional Schultheiss, affects linear S., generalized from S. R2 obtaining Nooten, for Mountain method Soutpansberg simple western and general the models. A of mixed-effects (2013). ants H. Schielzeth, epigeal & of S., inventory Nakagawa, An (2015). H. Africa. S. South Foord, Range, & C., T. Munyai, doi:10.1007/s10841-011-9449-9 ymclgclNw,29 News, Myrmecological 3,2023 doi:10.1016/j.tplants.2019.11.003 250-263. (3), fia ora fRne&Frg cec,31 Science, Forage & Range of Journal African eooi,84 Oecologia, idvriyadCnevto,21 Conservation, and Biodiversity ode 57 koedoe, ehd nEooyadEouin 4 Evolution, and Ecology in Methods ora fApidEooy 20 Ecology, Applied of Journal 3,3633 doi:10.1016/j.actao.2005.12.001 316-323. (3), 482-490. , 77.doi:10.25849/myrmecol.news 67-77. , 6,19-18 doi:10.1111/j.1365-2656.2008.01450.x 1191-1198. (6), 1,1-12. (1), uta clg,33 Ecology, Austral iest n itiuin,18 Distributions, and Diversity iest n itiuin,18 Distributions, and Diversity 9 3,2829 doi:10.1111/j.1442-9993.2007.01815.x 278-289. (3), 833-847. , 3,7789 doi:10.1007/s10531-011-0221-y 797-809. (3), 2,133-142. (2), 2,6-8 doi:10.2989/10220119.2014.939996 67-88. (2), ora fToia clg,31 Ecology, Tropical of Journal ple ymclg:awrdperspective world a myrmecology: Applied clgclEtmlg,43 Entomology, Ecological 029:067 5,4353 doi:10.1111/j.1472- 493-503. (5), 4,3040 doi:10.1111/j.1472- 390-400. (4), e htlgs,220 Phytologist, New caOecologica-International Acta rnsi Plant in Trends 3,379-388. (3), 267-280. , 1,10-24. (1), . Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. o n seblei Hluhluwe in assembalge ant for 2 Table species) of number (Effective Simpsons species) of AIC number (Effective Shannon model Best Richness Species Model Simpson and (Hy- Shannon diversity. communities numebers Simpsons effective ant and there Shannon affect by richness, represented complexity species are of habitat indices (GLMM) and Models size Effects Mixed forest. Tree rain Bornean 1: (2016). review. of Table canopy a A. high grasslands: W. the temperate in Foster, north Formicidae) & in menoptera: ants M., of K. role Yusah, The (2018). A. D. Landis, proce- 186 & D., bootstrap and B. location ”model-free” confound for Wills, analyses multivariate PIT-trap-A package Distance-based (2012). The R Y. responses. Wang, effects. & multivariate dispersion an T., (2017). S. discrete, Wright, mvabund- I., A. with D. Warton, Y. models Wang, (2012). regression & about I. doi:10.1371/journal.pone.0181790 L., inference D. for Thibaut, Warton, dure I., & D. Warton, T., S. data. abundance Wright, is multivariate doi:10.1111/j.2041-210X.2012.00190.x encroachment of U., ML. woody analysis Rosenzweig, Savanna Naumann, model-based - time Y., and (2017). space G. Wang, in Species-diversity Durigan, (1995). & J. P., G. Vermeij, B. Murphy, continents. E., three across C. widespread Lehmann, N., Stevens, 2,3338 doi:10.1007/s00442-017-4007-0 323-338. (2), odtoa Lklho ai n t infiac)admria ffcs(eine fpredictors of (Deviance) effects marginal and significance) its and ratio (Likelihood Conditional : etmdl n h agnladcniinlvrainepandbsdo eeaie Linear Generalized on based explained variation conditional and marginal the and models Best ehd nEooyadEouin 3 Evolution, and Ecology in Methods e equation: model Best Season + ˜Habitat** e equation: model Best Season + ˜Habitat** e equation: model Best Season*** + ˜Habitat* 03pn-0.19Wet - -0.37pen 0.15Wet - -0.3Open 0.21Wet + 0.15Open lblcag ilg,23 biology, change Global 6. . .80.27 0.08 0.22 1.1 0.23 0.12 364.9 0.11 1.7 417.2 1.3 549.84 10 ehd nEooyadEouin 3 Evolution, and Ecology in Methods 1,8-0.doi:10.1111/j.2041-210X.2011.00127.x 89-101. (1), et R best) Δ 1,235-244. (1), I (second AIC ymclgclNw,23 News, Myrmecological cec,270 Science, 2 m LSOE 12 ONE, PLoS 53) 503-504. (5235), 15-23. , 3,471-474. (3), Oecologia, R 2 c (7). Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. Str aia 53* 1503.8 705.1 1486.6 15.38** 26.2** 16.01** Soil Habitat Season Site Predictor Predictor c 00 1579.9 10.07 pc1 c .91491.0 9.99 pc1 ai L)Mria ffcsDeviance effects Marginal Deviance effects Marginal (LR) ratio Likelihood effects Conditional (LR) ratio Likelihood effects Conditional 11 Posted on Authorea 25 Nov 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160630390.02526653/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. 12