Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. etesse ao oGeraniaceae. to taxon sister the al. be et Aedo 1973; Yeo 1; (Fig. 2002). world 1998, the over 2002). all distributed al. et (Aldasoro nium America while El- North 2000), 1972; western al. (Guittonneau habits et area Aldasoro 1989; Mediterranean Oqlah, the in versity 97 lnsadGa 01.Mn ebr fGeraniaceae of members 1988, Many 2001). (Boesewinkel Grau Andes and central Slanis the 1997; of habitats subalpine Albers 1983; 1979, Venter 1979; b). Moffett 1996a, 1968; (Kers Asia n amr(93 showed (1993) Palmer and using Geraniaceae of study logenetic 2005), 2004, and 2000, 2006), 1999, 1998, al. et (Bakker eea n rsa aCna(akre l 98 99 2005). of 1999, species 1998, al. 39 et The (Bakker St. Cunha Madagascar, da Australia, Tristan Africa, and east Helena, in found number are lower Ka- species a of while Succulent KwaZulu-Natal Region, and Karoo, Floristic Nama Cape roo, the in centered Africa. species Southern in represented 2006). al. et Fiz 2002; al. et by (Aldasoro formed subclade the of group sister Geranium 2001). Grau car- and and Slanis fruit 1969; as (Hutchinson sepa- such are structure they features pel but morphological 2003) some II by APG rated 1993; Palmer and Devi Price Rama Hypseocharitaceae 1991; 1988; studies (Boesewinkel family Geraniaceae morphological family monogeneric the within and II the molecular APG included of 2000; have al. number et A (Soltis 2003). Ledocarpaceae, Vivianiaceae Greyiaceae, and Francoaceae, Melianthaceae families the with 07.Acrigt rc n amr(1993), Palmer and Price Geranium and to ITS According on 2007). data by ported of inclusion The ceae. Pelargonium sonia oyih 08b h mrcnSceyo ln Taxonomists Plant of Society American the by 2008 Copyright © Botany Systematic hlgn n itrclBoegah fGrnaeei eaint Clim to Relation in Geraniaceae of Biogeography Historical and Phylogeny rc n amr(93 rpsdthat proposed (1993) Palmer and Price eaica aeawrdiedsrbto u r best are but distribution worldwide a have Geraniaceae hlgntcrltosiswr xmndi orgenera: four in examined were relationships Phylogenetic along Geraniales order the in included are Geraniaceae The uoaycudesr ufcetse rdcinadsurvival. th and in production habitats seed disturbed often sufficient and ensure evoluti temperate could the cold, autogamy summary, to In Africa ranges. Southern mountain from American South in and hemisphere ra.A netro h ags ld fGrnaee( Geraniaceae of clade largest the of coinciding ancestor Miocene, An the s areas. in during Oligocene, occurred the diversification during their occurred However, probably Geraniaceae of branches monophyle stem are Geraniaceae by and formed Hypseocharitaceae Geraniaceae, data these to According stelreto h aiy opiigaot40species 420 about comprising family, the of largest the is Abr 96,19b lesadLbet19) and 1996), Löbbert and Albers 1996b; 1996a, (Albers Keywords— Abstract— mrFz al ags aiaAacn alsAd,Js usGrí,and García, Luis José Aedo, Carlos Alarcón, Marisa Vargas, Pablo Fiz, Omar PieadPle 93 a ta.1997), al. et Pax 1993; Palmer and (Price r hlgntclycoe and close, phylogenetically are Erodium Monsonia 20) 32:p.326–342 pp. 33(2): (2008), ob itrt h te orgnr fGerania- of genera four other the to sister be to hools ( Chloroplast uooptblt,dseslvcrac,dogttlrne molecu drought-tolerance, dispersal-vicariance, autocompatibility, Monsonia a 4seisadsosisgets di- greatest its shows and species 74 has Tuomndue l 07.Tephy- The 2007). al. et (Touloumenidou elJrí oaiod ard SC lz eMrlo2 81 ard Spa Madrid, 28014 2, Murillo de Plaza CSIC, Madrid, de Botanico Jardín Real Sarcocaulon Geranium trnL–F Sarcocaulon nai fiaadsouthwestern and Africa inhabit trnL–F , and Pelargonium Erodium in 1 uhrfrcrepnec ([email protected]) correspondence for Author Monsonia rbcL rbcL Tuomndue al. et (Touloumenidou Hypseocharis ob nlddin included be to Erodium eune eeue orcntuttepyoeyo eaica n Hyps and Geraniaceae of phylogeny the reconstruct to used were sequences ) California and eune yPrice by sequences Hypseocharis California Erodium omnctn dtr akP Simmons P. Mark Editor: Communicating California a bu 270 about has n olnto Ecology Pollination and a losup- also was and Erodium Pelargonium ol ea be could rw in grows Geranium Fze al. et (Fiz Geranium codn omlclrdtn n ipra-iaineaayi,tes the analysis, dispersal-vicariance and dating molecular to According . nyin- only could Mon- Gera- and , Erodium, 326 ihtebgnigo ao rdfcto vnsi hi distribution their in events aridification major of beginning the with bu h lwrvstrrltosisi Geraniaceae. in knowledge relationships the flower-visitor complete the to about necessary is information further le,be,btefisadbrs olntr r eswell less are Pollinators birds. for and known butterflies bees, flies, Other in visiting 2006). frequent flies relatively is long-proboscid and largonium by nectaries deep is with Africa flowers syn- South pollination characteristic in most dromes the and of types One pollination shape. between flower relationships (2005) possible of al. the evolution et ing the Bakker of 2002). analysis Kandori an 2000; published al. et Dlusskii 1989; and 1997) Struck 1997; Geranium Goldblatt and Manning 1996; Walt for der known are Geraniaceae the insects. short-tongued to accessible ily oge olntr,teetra nb()o h remaining the of con- (4) a knob of long- in species external to nectar the accessible the only pollinators, structure store tongued bag-shaped 1–3 or types tube 2007). of cealed al. nectaries et Touloumenidou the 2002; While 2001, 2000, al. et Aldasoro while flowers morphic tmn termiigseisof species nium of remaining base the (the at located stamens nectaries, nectar of knob-like closed external, species five forming (five 4) enlargement and stamens, sepal the to of due pockets base of the species at (two nectaries hypanthium hypanthi- the the in in tubes of tube a species by (most formed um nectary one of 1) nectaries types: five or four one have flowers Geraniaceae gomorphic. n rbadrn h n fteTrir Kr 1968; (Kers Tertiary the of end Africa the in during environments, extended Arabia steppe more and and progressively desert associ- became be to which can adaptation genera (Hutchin- with these of mass ated differentiation land The Afro-Arabian 1969). the son of characteristic are oemscevrnet Yo18,19;Boesewinkel 1990; 1984, occupy (Yeo and 1988). environments continents, mesic temperate northern more character- are today’s family of the of istic members other The 1983). Venter uhr fiao nsuhr fiapu h eierna area. Mediterranean the plus Africa southern in or Africa outhern eto ol,weeol eeaitpliainadfacultative and pollination generalist only where world, of rest e h anpliaosadohrrpoutv etrsof features reproductive other and pollinators main The Monsonia and tic. , no h eaica smre ytedseslo ancestors of dispersal the by marked is Geraniaceae the of on California Pelargonium a aig etre,POindexes. P/O nectaries, dating, lar California Gen17;Mlay18;Piip18;Hessing 1985; Philipp 1983; Mulcahy 1978; (Green osna Geranium Monsonia, Glbate l 95 odlt n Manning and Goldblatt 1995; al. et (Goldblatt , Erodium Geranium Pelargonium ooie ubro aiasi h northern the in habitats of number a colonised ) and and Smne l 00 and 2000) al. et (Simon Monsonia Erodium Pelargonium) , Pelargonium Erodium, ydoe r yshort-proboscid by are syndromes r itr otelretcaeof clade largest the to sisters are Vgl15,19;Ln 1990; Link 1998; 1954, (Vogel ) , unJs Aldasoro José Juan California, Pelargonium and )fv etre forming nectaries five 2) ; lwr r eeal zy- generally are flowers in Monsonia, California Pelargonium, and t Changes ate Monsonia Src n Van and (Struck eocharitaceae. Monsonia Erodium n all and lto the of plit eractino- bear Monsonia) ;3 five 3) ); 1 Thus, . study- seas- is Gera- Pe- , Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rdadtit n N eune eeicue nteaayi,of analysis, the in included were sequences DNA 44 one which thirty locations and three dred or two laboratory. in the identifi but in collected, measured time, were and Insects one 3). Appendix only visiting 2005; censused of (December-January, difficulty were the they to Due tats, 3). Appendix in specified ie My uut n etme,2004 September, and August, (May, times fiaadMdgsa;E utai,Tsai n e eln;F: Europe. Zealand; H: Mediterranea Anatolia; New G: and Islands; and Arabia Pacific eastern region, Tasmania and D: Indonesia Australia, Africa; Indochina, Asia, E: Southern eastern C: Madagascar; America; and South Africa B: America; Central and Geranium yeterl fteefaue nrlto omi evolutionary main to events. relation biogeographic in and features ana- these to of serve role would the lyse numbers chromosome about. and disper- and systems came reproductive sive habitats, distributions about data geographic of Compilation current in the ways the which examine and Geraniaceae of reconstructions netic fvst e ie n h ubro aao olntr eercre in recorded were pollinators of of taxa Species of table. number the a number and Absolute time, 3). per (Appendix visits field of the in observation by pollination. obtained similar were free A for counted. the tested mericarps were and mature plants opening, the unbagged before of in bagged number produced inflorescences were seeds plants Twenty 2) of four self-pollination. Appendix number or automatic (see three for Garden to test Botanical belonging Madrid to the used female litera- (Cruden in were the species per grown each from male of ovules Plants system data reproductive ten fertile 1977). the and define of least observations to used number field a were ture, plus (the of results, five number These by grain flower). divided For pollen species. was mean each flowers for the examined species, were plants dioecious different from were taken pollen 3 Buds flowers all at and five. counted microscope by dissecting were a grains divided under opened and were in Anthers flower 1989). preserved per and anthers multiplied fixed counted, of collected, P/O was number calculating anther for per the Thus, grains by carpel. pollen each of in number the one flow- indexes, ovules, Geraniaceae fertile 2). five (Appendix have species ers many for calculated were Real from dexes at obtained housed are field, (MA). Vouchers the Madrid bought. de in cases Botanico two collected Jardin speci- in seeds Living or gardens, from 2). botanical (Appendix obtained examined were were mens livin specimens studies, herbarium pollen and specimens and reproductive herbarium morphological, and For fresh 1). included (Appendix extractions DNA for used terial 327 GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ 2008] ra sdfrDV nlss h nesso h ubro endemic C: of Genera: number area. the each show in insets genus The each for analysis. species DIVA for used areas C mlfcto,Sqecn,adSqec Analyses Sequence and Sequencing, Amplification, PCR otdt bu lwrvstr eecmie rmltrtr;terest the literature; from compiled were visitors flower about data Most in- (P/O) pollen-ovule systems, breeding on information obtain To Features Reproductive of Analyses and Material Plant F h i ftepeetwr a oudraephyloge- undertake to was work present the of aim The IG .Dsrbto a fGrnaeeadHpecaiaeei the in Hypseocharitaceae and Geraniaceae of map Distribution 1. . ,H: trnL Hypseocharis – F n 21 and Geranium M rbcL TRASAND ATERIALS × ,M: anfcto.Toatesfo aho five of each from anthers Two magnification. eune eeotie uigti project this during obtained were sequences and Monsonia “ Erodium e liquid Kew ,P: M ETHODS – eeuulycnue three censused usually were Pelargonium 06 oa ieo censuses of time total 2006, ” California FrmnadBridson, and (Fordman ra:A North A: Areas: . — ,E: Monsonia h ln ma- plant The — Erodium n hun- One habi- ,G: ed n g 02 n utdvie(e.0.50,Ton n ihn 02.A50% A 2002). Kishino and Thorne 09.25.03, Sanderson (ver. (http://ginger.ucdavis.edu/r8s/; wer Multidivtime reconstructions and 1.70 These 2002) ver. 2002). (BRC) 1998, r8s Clock al. by et Relaxed performed Thorne Bayesian 2001; the al. et and (Kishino 2002) Likelihoo (Sanderson Penalized the (PL) marker: approach each for times divergence reconstruct to purpose. demands this program t for from The selected obtained 1). was likelihood highest analysis (Fig. the Bayesian with taxon tree endemic the the trees; one resolved by fully least defined were at endemism of of presence Areas 1997). Ronquist (DIVA; analysis eercntutdfrboth for reconstructed were character of distribution the map (Mad- to 3.08 changes. used ver. state was MacClade 1992) stationarity. Maddison reaching and to dison prior burn-in as tions nium 2Rad16Rfor 1460R and 724R forward DNeasy The the Germany). using Hilden, extracted Laboratories, primer (QIAGEN DNA Kit the Mini and Plant taken were material ( barium outgroups were which Viviania, of Melianthus, seven Greyia, Francoa 1), soma, (Table taxa 46 ing ( analy- phylogenetic the in of included ses were taxa Eighty-five included. were Two 1). (Table ec mn eune a acltdb sn h Jmto (Saitou 1980). method (Kimura model NJ distance the 2-parameter Kimura using the by with diver- 1987) calculated Nei Pairwise was and Jersey). sequences New among Branchburg, gence Biosystems, (Applied program se- (2002). were al. et products Fiz Cleaned to kit, according California). Clean-up ampli quenced (PCR Carsbad, The columns respectively. Laboratories, filter (2000a), spin al. MoBio using et purified Savolainen were and products (2006) fied al. et Fiz of in amplification the for conditions PCR The trnL unn h asmn ace Nxn19) re n aamtie for matrices data and Trees 1999). rbcL (Nixon ratchet Thu parsimony replicate. the (M first running PRAP/PAUP the using in them aborted analyzed analyses we the data Since large, sequenced. very are not matrices parts the for marks value ftesqecs olwdb aulajsmn omxmz similarity maximize to adjustment 2004). manual (Simmons, by followed sequences, the of Geraniales. in 18.22% and Geraniaceae For and Geraniales. Geraniaceae in in 27.94% 45.89% reached taxa between sequence in divergences rtosuigtedfutprmtr.A5%mjrt uete a ob- was tree 10 rule gen- first majority million the 50% 2 discarding A for after parameters. run tained default was the 2001) using Ronquist, erations and (Huelsenbeck 3.0b4 ver. in (13 18% ones informative the and subset). Geraniaceae), Geraniaceae in (19% Geraniales in the selected tests Both + GTR 2002). models: employ- (Nylander substitution used same software an were 1.1b 1974) 1988) Akaike MrModeltest Felsenstein (AIC, ing (hLRT, criterion test information ratio Akaike the likelihood hierarchical the data, and Nixon 1989; (Kluge, the combined analysis we no 1996) simultaneous and Carpenter replicate For per replicates swapping. 10 1000 branch with using addition se- performed taxon option were random Mulpars 1985) replicates, (Felsenstein per- the analyses with was Bootstrap and lected. search swapping, Bisection- Tree branch heuristic sequences, (TBR) addition A random Reconnection of 2003). replicates ver. 100 MrBayes Huelsenbeck with and formed and 2002) (Swofford (Ronquist 4.0b10 PAUP* 3.0b4 using out carried were the ils(4 nGrnaeesbe) rmwihifraieoe a 33% For was Geraniaceae). ones informative in which (30% from subset), Geraniaceae in (44% niales for I optimisation. ACCTRAN charac- using weighted performed were equally using searches unordered, conducted Heuristic were with ters. analyses 1971) Cladistic (Fitch analysis. parsimony this Fitch in used not were Francoa w eae-lc ehd sn ifrn ttsia ae eeused were bases statistical different using methods relaxed-clock Two igorpia nlssadMlclrDatations Molecular and Analyses Biogeographical prxmtl 0m fla isefo rs fedcletd rher- or collected) (field fresh from tissue leaf of mg 20 Approximately The Seqed the using edited and file contig a in stored were data Sequence lsa e.16b(hmsne l 97 a sdfrtealignment the for used was 1997) al. et (Thompson 1.62b ver. X Clustal aeinMM nlss(agadRnaa19)uigMrBayes using 1997) Rannala and (Yang analysis MCMC Bayesian A the fit best that evolution sequence of model simplest the determine To hlgntcrcntutoso the of reconstructions Phylogenetic trnL (UAA)- and two , rbcL trnL trnL ‘ – e and F ’ h rprino aibestsfor sites variable of proportion The . trnL – n ees primer reverse and arxadi a o osbet lg hm osqetythey consequently them; align to possible not was it and matrix – Hypseocharis F trnF F Crossosoma Tbe1 pedx1 i.1,too hc eeoutgroups were which of two 1), Fig. 1; Appendix 1; (Table – eune a 217 had sequences F aionamacrophylla California GA pcr(aelte l 91,adpies1,636F, 1F, primers and 1991), al. et (Taberlet spacer (GAA) r otda reAE(td ubrS1569). number (study TreeBASE at posted are Balbisia rbcL ,16 .Aayi of Analysis ). eune Omta ta.19;Fye l 1998). al. et Fay 1992; al. et (Olmstead sequences and rbcL rbcL Monsonia trnL Wendtia ‘ h rprino aibestsws26% was sites variable of proportion the , f eune,dfeecsrahd1.5 in 11.05% reached differences sequences, ’ – eeue o C mlfcto fthe of amplification PCR for used were – ⌫ trnL 1 pdpnigo aa h largest The taxa. on depending bp 413 4 F o the for ( and eune ls40 plus sequences rbcL ,12 rbcL – rbcL hwdn dnia rget in fragments identical no showed F and rbc , arx r10 or matrix) Pelargonium ü and trnL eune a efre us- performed was sequences trnL lr20,20) hc allows which 2005), 2003, ller trnL sn dispersal-vicariance using L rbcL – Wendtia – trnL – F F F n obndmatrices combined and aaes sn missing using datasets, arx n T + GTR and matrix, and trnL – F n 1outgroups 11 and ). a 0 nGera- in 50% was rbcL 5 — – Erodium F ( esm,Crosso- Bersama, trnL neta areas Ancestral n combined and r described are – F ,48 genera- ) Gera- ⌫ he s, d d + e - Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 328

TABLE 1. Dates of nodes of Geraniaceae tree in MYA. * Indicates age constrained in this node.

Separation of Separation of Separation of Separation of Monsonia and the Separation of Separation Geranium two main groups of Separation of Pelargonium and ancestor of California and the and the ancestor Separation of Separation of subgenera of Geranium: two main subgenera Separation of Separation of the rest of California, Erodium ancestor of Erodium of Erodium and Erodium and California, and Robertium and Erodioidea of Erodium: Malacoidea basal-most branchs basal-most branchs Nodes Geraniaceae (D) and Geranium (F) and Geranium (N) California (H) Geranium (O) Erodium (L) plus Geranium (K) and Erodium (L) of Monsonia (G) of Pelargonium (E) rbcL r8s-PL 38–47* 28.20 19.94 — 18.0 — 14.46 15.46 14.52 11.47 rbcL Multidivtime (s.d.) 38–47* 26.93 (3.7) 22.4 (3.61) — 20.64 (1.2) — 18.14 (3.49) 17.8 (3.52) 17.5 (4.58) 14.18 (3.45) trnL–F r8s-PL 38–47* 32.71 — 21.73 — 20.34 15.57 16.33 13.89 23.00 trnL–F Multidivtime (s.d.) 38–47* 33.82 (3.18) — 28.57 (3.69) — 27.31 (3.7) 23.82 (4.1) 24.36 (3.98) 22.47 (4.57) 24.28 (5.31) ettv,bcuete eepoal ipre ytemnt different to to with respect man marked the with species by 2 dispersed of probably continents. distribution Fig. were geographic they in The because each differently are tentative, 4. topologies of marked and the distribution 3 are As Figs. interpretation). the nodes area represent the for 1 right different, inter- Fig. the area (see for analysed on 1 taxon Fig. letters (see parsimony-uninformative analysis The lett DIVA (excluding non-circled pretation). from names; 0.637 inferred node areas are = ancestral letters are CI Circled 0.884. 683, = that num- RI was The tree characters), branch. steps the parsimony under of * the by ber in indicated are Branches tree Bayesian probabilities. contradicted posterior are below tansfrtotria ucae eetknfo h oslrecord fossil the from con- taken age were as Minimum data subclades D. these used terminal node we two Thus, for 2005). for constraints al. straints et maximum Bakker also and for see D minimum but MYA (node 3, 38 Hypseocharitaceae 2, and excluding Figs. 47 Geraniaceae in between Fagales of ranged the node authors of ancestral fossil these the a by with Wik- obtained it calibrating ages of by The data tree angiosperm the the from of clades deduced was constraint str age upper An test. The for analyses. ratio marker. used likelihood each the was for using tree) node rejected each BI was hypothesis of the clock times from divergence (obtained the tree reconstructing consensus rule majority in airto onswr bandfo osl n ulse dating published and fossils from obtained were points Calibration F ö trnL IG ta.(01 pedcs oe13,wocluae h g fmany of age the calculated who 183), node appendices: (2001, al. et m .Src osnu fte192ms asmnostesobtained trees parsimonious most 1,962 the of consensus Strict 2. . – F nlss ubr bv rnhsaebosrpvle;those values; bootstrap are branches above Numbers analysis. ● ers is . Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. oe n pe ii f0ad09(Magall 0.9 and 0 of limit upper and lower ae,asmn httert frltv extinction calcu- relative were of diversification rate of the rate that of assuming lated, Estimators (2001). Sanderson and data. estbranches the of form values default obtained were values 0.4 Multidivtime. other MYA, package of 100 at value set was value a Bigtime MYA selected 45 we equals which deviation 1.5, of value a prior) (rttm used (1.5 root we the deviation, of standard age the its for and procedure were: Multidivtime Bayesian in a dis used using settings posterior times The node the using and estimate rates estimated substitution to tributions, were served Multidivtime Baseml, covariances) using Finally, and data Estbranches. the variances from (and estimated were branch-lengths parameters the 1998) After al. 2002). et Baseml 9 Thorne programs: (ver. 8-5-03; (ver. Multidivtime three Estbranches and of 1997), Yang use 3.14; the ver. (PAML involves which (2004), Rutschmann data validation the cross for model lowest semiparametric (the best 2002). the error (Sanderson provided prediction therefore it lowest score); the to sponded 0) triga n nraigtevleo yseso . pt 1,012. to up 0.5 and of steps of by x value of value smoothing the increasing The and 1 at starting 10x), aiainsoewsfudb etn ifrn mohn aus( values smoothing different testing by found was score validation mndb h uhr n w fte eeasge to assigned ex- and were were them grains of these two of Barbata and authors Photographs the 1989). Mio- by Campo middle amined (Van the from Spain grains of pollen cene are Geraniaceae to assignable dently arw f (arrows 329 GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ 2008] 7ad3 Y.Tedtslne yarw ondsaemnmmaeconstraint age minimum are nodes to arrows by dis linked inferred dates indicate The arrows MYA. by 38 linked and letters Circle of 47 0.783. pairs The = analysis. RI in DIVA those and values; characters), bootstrap are parsimony-uninformative parentheses (excluding in not are that branches beside n etre pt he ie oaodlclsaiiy h smoothing The stability. local avoid to times three (log parameter to perturbe was up estimate Searches Each restarted estimates. 2002). and time (Sanderson different five used at started was were (PL) method likelihood 1991). penalized Blackmore and Stafford see elements; supratectal of by rest covered the in ticulate; utdviewsue olwn h rcdr ulndby outlined procedure the following used was Multidivtime ae fdvriiainwr banduigeuto fMagall of 7 equation using obtained were diversification of Rates F oetmt h iegnetmso h eaica oe,ter8s the nodes, Geraniaceae the of times divergence the estimate To IG × Geranium .Crnga eosrce fe analysing after reconstructed Chronogram 3. . 0;frtepiro aecag bonen.Frisstandard its For (brownmean). change rate on prior the for 30); ple ihitroe tie h eto ugnr sreticulate), is subgenera of rest the striae; interwoven with (pollen 1 n f and subg. 10 2 nFg ) h nyaalbeGrnaeefsisconfi- fossils Geraniaceae available only The 3). Fig. in ␭ a acltdb rs-aiain h oetcross lowest The cross-validation. by calculated was ) Robertium Geranium – 25 – 3 ihn ta.20;Ton n Kishino and Thorne 2001; al. et Kishino 03; ␭ ple ihsxn itnt eryre- nearly distinct, 2 sexine with (pollen =32( rustesxn sidsic,adis and indistinct, is sexine the groups rbcL and ) ó n adro 2001). Sanderson and n ␭ =10( ∈ rbc aidbtena between varied ihrs twsotie rmacnesso h Psotie in obtained MPTs 6 the of consensus a from obtained was It r8s. with L trnL Erodium – F corre- ) subg. ␭ ó d = n - . etr r oenms o-ice etr r neta ra inferre areas ancestral are letters non-circled names; node are letters d aetee r otro rbblte.Tenme fsesws76 I= CI 776, was steps of number The probabilities. posterior are parentheses esleet.Nd a osrie o iiu n aiu ae of dates maximum and minimum for constrained was D Node events. persal i.2,adteohrfre ytermiigsbeea(K subgenera remaining the by formed other the and 2), Fig. aeinaaye Fg.2 main (Figs. Two analyses Bayesian analyses. all the across dium in consistent appeared is branches This 2). Fig. n hwterslsotie rmthe from obtained results the show 3 and 2 eedn aast u ifrn nisbac support branch its in in- differing the of 4). but that (Fig. sets to similar com- data topology The dependent a Hypseocharitaceae. produced plus matrix Geraniaceae bined to the sister However, 2). as Fig. D, (node alone rbcL Geraniaceae of and that C) for (node clade Hypseocharitaceae the plus Geraniaceae for by support analysis formed high with Bayesian topologies, similar and produced Parsimony respectively. sequences, nd for H comprising (node clade large a Monsonia nium 1) rnhs n omdb otseiso subg. of species most by formed one branches, 3). 2, in- (Figs. topologies markers alternative the both for from support ferred low the of because relation- phylogenetic between The ships polytomies. nested used. two locus showing the on of clade depending The topologies different showed nlsso trnL of Analysis The h ld ftegenus the of clade The Hypseocharis f8MAfrtotria ons(rosf (arrows points terminal two for MYA 8 of s n h oohl fti eu swl upre in supported well is genus this of monophyly the and , reidctslwspotfrtermiigGeraniales remaining the for support low indicates tree ssse oaltermiigGrnaee(oeD,3) D), (node Geraniaceae remaining the all to sister as trnL nldn h section the including , – F Monsonia trnL and eaim Erodium, Geranium, ssse oGrnaee(oeC,2) C), (node Geraniaceae to sister as – rbcL F – ,rc n obndMatrices Combined and rbcL F, n oeNfor N node and a neovdwt epc to respect with unresolved was re hwdfu annse clades: nested main four showed trees – Geranium R 4). trnL ESULTS Geranium – Sarcocaulon F and and , Erodium, rbcL California rbcL hwdtomain two showed rbc .Tefut clade fourth The ). 1 nlss Numbers analysis. L nd ) n 4) and G), (node n f and nlssof analysis trnL Robertium and 2 – ). r unclear are F — California and Pelargo- Figures trnL-F from d 0.517 rbcL Ero- (K 1 2 , , ; Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. niae y*udrtebac.Tenme fsesws149 I=066(ex 0.656 = CI 1,439, was steps of number The branch. the neau under * by indicated onamcohla rdu rcyapm .glaucophyllum, E. macrorrhizum, brachycarpum, of Geranium positions Erodium the macrophylla, only analysis, fornia simultaneous the In 3). 2, the both for similar quite were 3 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 330 aetee r otta aus hs nprnhssaepseirpro posterior are parentheses in those values; bootstrap are parentheses F h oooisotie yprioyadB analyses BI and parsimony by obtained topologies The IG ’ .Src osnu fte40ms asmnostesotie rmcomb from obtained trees parsimonious most 400 the of consensus Strict 4. . 19)casfcto of classification (1990) s n h rnhof branch the and Erodium trnL n Yeo and – F and ’ 19)casfcto of classification (1990) s rbcL osnanivea-M. Monsonia arcs(Figs. matrices Cali- aiiis rnhsi h asmn rewihcnrdce aeint Bayesian contradicted which tree parsimony the in Branches babilities. Geranium heliotropioides ofit nthe In of conflict. position the only compared, are Geranium in while ldn asmn-nnomtv hrces,adR .4.Guitton- 0.841. = RI and characters), parsimony-uninformative cluding r niae ntergtsd ftetree. the of side right the on indicated are ined rbcL ete oooyhdsrn otta support. bootstrap strong had topology neither ; rbc and L ti itrt h ld omdby formed clade the to sister is it r ncnlc Fg ) If 4). (Fig. conflict in are trnL trnL-F – F tree, ubr eiebace htaentin not are that branches beside Numbers . California aionamacrophylla California trnL ssse to sister is – F and Erodium rbcL Erodium e are ree trees sin is and , Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. Erodium, in e ieprya ssy for (s/s/y) year per site per tions raniaceae. of clade Geranium largest the of nium diversification the Also, wards. lower the dium to of Oligocene diversification late The the Miocene. from range ages These by MYA. formed clade the the of Erodium, separation re- Africa, of our age in to the According distributed branches sults, region. Asian these be West between and should Mediterranean, splitting which late of the MYA), indicate time (26–34 nodes the these as for obtained Oligocene data The 1). Table 3. Fig. of for isof cies (0.1–0.23). rates lowest to the contrast showed in rates high show oa ubro pce 1 0 203 4074 c420 39 15.4 14.96 c270 14.52 801 11.47 810 45.8 64.32 species autogamous mostly or facultative of rate Pergentage extiction for rate rate Diversification extiction (MYA) genus for the rate of Diversification split for proposed Date species of number Total 331 GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ 2008] eimt o / nie Fg ) hyaegnrlyau- generally are They 5). (Fig. indices P/O low to medium otenArc c.Tu,adsesleettwrsthe towards event ances- of an dispersal for tor required a was region Thus, Asian Mediterranean-West (c). of Africa that while the Southern F), in node cg, region (area Asian Mediterranean-West plus and of nested ancestor The the c). by (area formed Africa clade Southern to assigned was D) (node Hypseocharitaceae plus Geraniaceae C). the (node for as suggested area was South bc) nodes). ancestral (area the Africa over Southern letters plus see America respectively, re- 3 2, reconstruction (Figs. the optimal tree for single dispersals a 14 in quiring resulted DIVA using utai) hc sepandb ogdsac dispersal plus long-distance (Mediterranean a eg by area explained event. is to which assigned Australia), was for which except region, one Asian Mediterranean-West of the nodes to lower assigned dis- the to Conversely, due probably events. areas, persal of combinations large to assigned rnhs hs aayeddart f86 10 * 8.64 terminal of the rate a constrain yielded to data These used branches. were Wikstr of fossils data genus Geraniaceae the the of from split tained the for constraints T cnieigPOadatmtcslig 49191. 2 37 c20 15.3 1.9 14.9 — selfing) automatic and P/O (considering al hw h iesfcto ae o h aiyGe- family the for rates diversification the shows 2 Table of ancestor the of separation the marks F Node / nee n olnto nGeraniaceae in Pollination and Indexes P/O igorpia Analyses Biogeographical h ld fteacso fconnd fGeraniaceae of node crown of ancestor the of clade The eosrcino iegneTimes Divergence of Reconstruction of nodes upper the of Many ABLE trnL-F California em ob eet uigteltrMoee( Miocene later the during recent, be to seems em ohv curdfo h i-icn on- mid-Miocene the from occurred have to seems osna Geranium Monsonia, Pelargonium .Dvriiainrtso eaica sn bLdt n oefeatures some and data rbcL using Geraniaceae of rates Diversification 2. and eut r hw nTbe1adtechronogram the and 1 Table in shown are Results . and es ao ruigmr hn30species) 380 than more grouping lato, sensu Pelargonium California a sindt h opst otenAfrica Southern composite the to assigned was California r itiue nArc Fg ) n have and 1), (Fig. Africa in distributed are Taxa 02–.2 and (0.28–0.42) n h netrof ancestor the and ndsNo )i ewe 18–29 between is H) or N (nodes , Erodium, — trnL Geranium ∈ ∈ Erodium osna Geranium, Monsonia, osna Geranium Monsonia, h igorpi analysis biogeographic The Pelargonium ö . .67004 .87014 .540.1336 0.2336 0.2564 0.3698 0.1045 0.2046 0.2857 0.4277 0.0948 0.1308 0.0677 0.0933 0.9 = 0 = ta.(01,wietwo while (2001), al. et m – F rbcL and read1 o the for 11 and tree Pelargonium n .610 2.26 and , eeursle and unresolved were Geranium and — California pe n lower and Upper Monsonia a sindto assigned was Monsonia Erodium — −10 trnL (0.25–0.37) . otspe- Most Geranium Geranium :Hypseocharitaceae B: , eeob- were substitu- −9 Erodium, and G lsGrnaeeC eaica E: Geraniaceae C: Geraniaceae plus – Fg 3; (Fig. subg. . F which s/s/y Gera- were rbcL Ero- tree , , e eu n h / ein band hyaetknfo biblio- from 2. taken Appendix are in They compiled obtained. data medians field studied P/O and species species graphic of the of numbers and number total genus the are per genera are the box-plots under the Numbers studied. over Numbers Geraniaceae. of onoptbeadvstdb o ubro visitors, of and Lepidoptera, number Diptera, low long-tongued a to medium by mainly visited and toincompatible htvstr r eaieysac in our scarce Although relatively genera. suggest are of they visitors limited, rest quite the that are time in per than visits on higher observations is it and 6), and species, of indexes P/O Lepi- The pollen. a for bees, of 11,753 lot by large a produce wasps, visited and etc.) (Bombylidae, are doptera, pollinators and of nectaries, range knob-like accessible, of species their have remaining and The indices P/O unknown. low are have pollinators Africa east in grow that to taries Two 5). limited Fig. 3, are Appendix observations our insects, long-tongued but by pollinated probably are and indexes compar- genus. are this in auto-pollination rare short- and atively birds, wasps, by Pollination Diptera, 3). tongued Appendix 3, (Table bees large esn h ubro iio aaprseisi ut similar quite is species flowering per in the taxa visitor of cases of end auto-pollination number the many The cases, at season. in or some situations are In stressful they in 5). increases and Fig. P/Os 2; low (Appendix selfers with species of tion 1). (Fig. habitats disturbed 32.87% places. all disturbed and of mountainous in grows it where Geraniaceae. for F Monsonia Erodium Erodium IG .Bxposo / nee nanasadprnil fgenera of perennials and annuals in indexes P/O of Box-plots 5. . eae otpso erdcin(:ntstudied). not (—: reproduction of types to related Erodium .speciosa M. sdsrbtdmil nteMdtraenbasin Mediterranean the in mainly distributed is and pce ihnca-okt aevr ihP/O high very have nectar-pockets with species Geranium pce r uoai nul htcolonize that annuals autogamic are species Geranium Pelargonium n ,3 for 7,337 and ihamda aaprseis Fig. species; per taxa 9 median a with ( Monsonia Erodium Erodium G: Monsonia ihamda 1tx per taxa 11 median a with pce ihtblrnec- tubular with species .ciliata M. hwdalrepropor- large a showed Monsonia .drudeana M. K Geranium : r uncommon are n compar- and Monsonia Tbe3, (Table L Erodium : 6 Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 3 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 332

TABLE 3. Total number of species in each genus (N) visited by insects of each functional group, and median of taxa of visitors per plant genus (m). Last row (total) shows the total number of Geraniaceae analysed for visitors and the median of taxa numbers. The six functional groups are classified according taxonomy, tongue length, and body size. Data are extracted from Appendix 3.

Pelargonium Monsonia Erodium Geranium

Geraniaceae genera Number of species of Number of species of Number of species of Number of species of Median of Pelargonium visited Median of Monsonia visited by Median of Erodium visited by Median of Geranium visited by Insect groups visitors taxa by insects of this group visitors taxa insects of this group visitors taxa insects of this group visitors taxa insects of this group Long-tongued flies (Tabanidae and Nemestrinidae) 1 23 0 0 0 0 0 0 Medium-tongued flies (Bombylidae) 1 11 1 2 1 8 0 0 Short-tongued flies (Asilidae, Calliphoridae, Conopidae, Empididae, Muscidae, Syrphidae, Tachinidae, and others) 0 1 0 1 3 13 3 16 Wasps and related taxa (Vespidae, Eumenidae, Sphecidae, Sapigidae, Chrysidae, Ichneumonidae, and others) 0 2 0 1 1 8 1 4 Small bees (Halictidae, Andrenidae and Collettidae) 0 0 0 1 2 11 2 16 Large bees (Anthophoridae, Megachilidae, Apidae and others) 1 16 0 2 2 10 3 17 Total 1 47 2 4 11 13 9 20 ypisadsm te itr eg oblde r better are for Bombylidae) pollinators (e.g. Diptera Conversely, other to self-pollination. some and causing contrast Syrphids often in stigma, the but, reach androecium in As the espe- period. insects, encircle female Halictidae the some during of cially movements caused chaotic self-pollination This the grains) prevent stigma. not by the pollen does from but far with hinders away the (still process them of keep anthers to beginning outwards the bend the period at fertile and female dichogamy, incomplete show oe easadd o aerno oeet niethe inside movements random make flower. not do and petals lower each; h 100 than 3) more Appendix during censused genus, each in for species v/m 0.133 and 0.130 spectively in abundant atively aaaetknfo ilorpi n il aacmie nAppen- in compiled data field and bibliographic from taken are Data i 3. dix iiatymr ndmn p=002 = z 0.002, = (p total demand the in more of nificantly the analysis showed Mann-Whitney taxa A Syrphid large flat- promising nectar. by shining of Syrphids spots, attract or quantities which color of droplets, (e.g. all nectar hairs), petals tened mimicking upper hairs the globular on structures attractive four the for (v/m) minute of per species visitors 0.025 were: medians h ubro ypi iisprtm nti ihri spot- in = higher z is 0.0006, unit = time (p per species visits ted Syrphid of number the ihattdsadlttds ny86 falstudied all of 8.6% Only latitudes. and perennials. altitudes mountain high small-flowered especially and is Also, P/O annuals 2); in Appendix 5; low (Fig. rates selfing high and type. either they of visits of flowers number to the made in difference Lepidoptera), significant no and showed Coleoptera, Hymenoptera, Diptera, other F huhmn pce of species many Though Certain IG .Bxposo h ubro iio aarcre nec genus. each in recorded taxa visitor of number the of Box-plots 6. . Geranium Erodium Monsonia Geranium Erodium stegnswt oeseisaatdto adapted species more with genus the is sect. Erodium tde cnue uig2 ) n re- and h), 20 during (censused studied pce rsn eimt o P/Os, low to medium present species Malacoidea eas hyhvradln nthe on land and hover they because Erodium − and Erodium .3 3.Ohriscs(e.g. insects Other 13). = n 3.43, Geranium and Erodium pte pce ob sig- be to species spotted E r rtnru,they protandrous, are sect. . Geranium − Apni ) The 3). (Appendix .8 6.Also, 26). = n 3.08, and Cicutaria Erodium Geranium ml bees small , they , show Gera- (15 Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. hstp ffoe Tbe3 pedx3). Appendix 3; pollinating (Table while successfully flower genus, difficulty of this have type in this flies pollinators and effective bees most small the are bees liuedsubdhbtt.Telrefoeso ayspecies many of of flowers large The habitats. disturbed altitude nium 08 I TA.PYOEY&BOEGAH FGRNAEE333 the of GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ 2008] nietehpnhu) h euto ftesaesfo 15 from deep stamens tube the of (a reduction one the to hypanthium), nectaries the five inside of reduction apomorphic the of flower, and zygomorphy the include plesiomorphic synapomorphies these of of However, re- Some mixture traits. (1993). first a Palmer was show and which species Price Geraniaceae, by of ported genera remaining central the the of meadows subalpine in Andes. survived that s.str. that suggest relatives, closest Hypseocharis its with differences phological monophyletic as 2 taxa (Figs. both al. of position for et The support groups. Soltis good by reported Wikstr studied and who b), was (2000a, al. Geraniales et Savolainen order (2000), the in Hypseochari- plus taceae Geraniaceae of position were alone The Geraniaceae corroborated. the Gerania- and of Hypseocharitaceae Palmer monophyly plus showing The ceae and s.str. (2003), Geraniaceae Price II of group APG of sister the conclusions and the most (1993) for confirm by congruent obtained and were trees clades analyses The Bayesian 4). and (Fig. parsimony analysis simultaneous the in of position rophylla the incongruence: of point one only ags eaica ld curdi otenArc and between Africa split early Southern The in Mediterranean. occurred the clade Geraniaceae largest dispersal. and/or vicariance by explained 1 be can (Figs. connections that transAtlantic Geraniales several the be show Geraniales of to of families range Africa home Southern ancestral and the America South suggest analyses new a for information sam- sufficient extensive acquire treatment. More taxonomic to 1989). needed El-Oqlah is 1984; pling Yeo 1972; genera neau classi- the taxonomic current within their fication reflected partly only analysis Geranium 5 to ean nla.Acrigt the trees, to analysis distinc- According the unclear. confirm remains analyses present of The tiveness 2007). al. and et ITS (Al- idou and morphology 2001) on al. et based dasoro analyses phylogenetic by gested Monsonia, et (Aldasoro rims 2001). without al. bristles meri- fruit the and of (fusiform), shape carps include characteristics plesiomorphic The vlto fteGeraniaceae the of Evolution codn oDV,sprto ftesse eeat the to genera sister the of separation DIVA, to According Estimates Age and Biogeography present the in retained groups infrageneric the Finally, genus The h rsn eut ao h nlso of inclusion the favor results present The Geranium – ,adaeohru eiap sitdb pappus. a by assisted mericarps anemochorous and 8, N=16 r uoai nul rwn nlow- in growing annuals autogamic are 186) = (N trnL – nadto,temi rnhswr elsupported well were branches main the addition, In . ) h ereo oeua iegne n h mor- the and divergence, molecular of degree the 4), and – spooe yAbr 19a 96)adsug- and 1996b) (1996a, Albers by proposed as F California satxnrltdt netr fGeraniaceae of ancestors to related taxon a is Pelargonium ao agrvstr.Lrebe n bumble- and bees Large visitors. larger favor and Erodium California rbcL lhuhispyoeei position phylogenetic its although , . eune aecnessteswith trees consensus gave sequences Hypseocharis ih esse ru ftecaeof clade the of group sister be might D a on ob h itrgopof group sister the be to found was ISCUSSION Geranium trnL — h hlgntcanalyses phylogenetic The – ntepyoeei trees phylogenetic the in F rbc and — akr (Touloumen- markers n simultaneous and L h biogeographic The Hypseocharis Erodium Pelargonium ö ta.(2001), al. et m Sarcocaulon aionamac- California Pelargonium (Guitton- – ) The 3). sthe as and in erna ai rws sat utai.As several Also Australia. to Asia Medi- of the west groups from or dispersal re- Basin distance Asian terranean long Mediterranean-west suggests and This Australia and gion. the region to Asian one Mediterranean-west just the to of clade assigned central were the to sister nodes Many vironments. of species current have and the some could of regions of mountain ancestors these the many in part encouraged changes of latter climate the appearance the and The systems during Pliocene. Eurasia and Mediter- in the Miocene and in mainly Basin occurred ranean diversification this species 3), 400 (Fig. than more with Geraniaceae, of of al. clade et largest the (Lavin America North to then and 2000). Europe North from to or 1969), Africa Bakker later Zinderen and (Van deserts America Saharo-Sindian North the to via Eurasia South- to from Africa dispersed have ern may Geraniaceae of of clade largest dispersal ancestors for a occurred have 4), California must (Fig. America North accepted to is event tree analysis simultaneous lms ws aps rahsv eie)se o to not seem devices) adhesive or pappus, awns, plumose 2006). hemisphere al. northern et Fiz the 2003; and (Smissen been Australia has between dispersal described long-distance Successful America. South et (Bakker Africa 2005). Southern 2004, in 1972; age, al. diversification (Axelrod this of During Valley rate 1972). greatest Rift al. and the continent et and of the Baker of development Miocene uplift the further late of the the result a during as largely Africa Pliocene, in accelerated greatly h egeacret(ise 90.Teporsiechange mid- progressive The the 1980). in (Siesser current place Benguela take the have in- 10 Zinderen to (around most Miocene (Van the seems MYA However, aridification 2004). 34 al. tense Oligocene, et early Dieckmann the 1975; Bakker to (Ward MYA 1983) 80 al. Cretaceous, et the since occurred Africa have may southwestern episodes in Arid Africa 1978). central Raven west rain- and present-day lowland of (Axelrod by those occupied to was similar 75 very Africa Between forest of semideserts. much and MYA, deserts 30 to and adapted best the are of species tolerant; commu- pollinator of probably composition midMiocene, the nities. in the changes during by Africa driven Southern knob-nectary in be- and sible splitting deep-nectary phylogeny, to the Geraniaceae convergent According tween of be clades. reconstruction also different our could in nectaries derived concealed synapomorphies case, character. derived that a be In may flies long-tongued by pollination of ancestors the iesfcto ftetomi ie fdogtaatdGe- drought-adapted of the lines triggered raniaceae: main have two might the climate of colder diversification and drier a towards akre l 20)rpre htteacsrlpollination ancestral the that reported of (2005) system However, al. split. already et early their were Bakker during Africa pollinators Southern long-tongued in present that suggest could ial,i sipratt omn ntedvriiainof diversification the on comment to important is it Finally, oeo h hrceitc fGrnaeemrcrs(e.g. mericarps Geraniaceae of characteristics the of Some genus the of position the If Most Geranium Geranium Pelargonium , Pelargonium Erodium, Pelargonium Geranium n 4of 74 and oaatt itre rt ihadcl en- cold and high to or disturbed to adapt to Monsonia Monsonia and – 5MA fe h ia salsmn of establishment final the after MYA) 15 emt aedsesdrpael to repeatedly dispersed have to seem and sb hr-oge le rbe,while bees, or flies short-tongued by is n h fleshy the and Erodium Geranium , rdu,Geranium, Erodium, Monsonia sects. California codn oorestimates our to According . hs h netr fthe of ancestors the Thus, . Sarcocaulon Pelargonium Monsonia pce r drought- are species nthe in Monsonia Aridification . and and rbcL ece its reached California Monsonia Erodium Erodium n the and sfea- is Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. titdt otenArc n aii.I otat north- contrast, of In Namibia. species and ern Africa Southern to stricted 33 [Volume BOTANY SYSTEMATIC of species many that gonium concluding capa- Geraniaceae, certain da dispersal in wind compared bility Tristan of authors indicative and Some parameters Helena, and 1998). characters al. St et Australia, (Bakker to Cunha Africa several South Also from 2006). al. et of (Fiz species America or Mediter- Australia the to from ranean events dispersal several experienced have certain DIVA that However, diversification indicated dispersal. area, analyses long-distance distribution for their chance nor of rate, extent the to relate 334 California and pollina- Most their taries habitats. on and depending ecology pressures tion selective subject be different could to Geraniaceae of groups main the distribution, animal herbaceousness. and and Also, diversity signif- pollination species a 1996). high of between (Johnson existence relationship the mosaic icant reported (2000) pollinator fauna Renner and a pollinator Ricklefs or of depauperate 2006) existence a Barraclough 1996; the 2006), Johnson 1992; al. Bond and et 2006; (Johnson Manning Niet and der Goldblatt Van 2006; in (Barraclough pollinators ex- region specialized alternative of the incidence three higher Manning proposed the and have planations: Goldblatt authors 2000; reported Various Steiner been 2006). and have Johnson sys- 3; insects, pollination (Table long-tongued specialised involving other tems, In the several pollination. long-tongued between Africa, to clades shift Southern a certain and species in of al. correlation number et a Bakker re- reported Also, (1996) syndromes. (2005) pollination Walt and der shape flower Van and that Struck ported 2001). Sanderson of and those to not nium, but 2), (Table sperms di- faster of a diversification to Monsonia of lead rates The can rate. and versification evolution during force selective Rates tion both in which observed endo-zoochory, been as or have such strategies ecto- events, and dispersal rare myrmecochory these Thus, sporadic by complexity obtained. their figure increase could final influ- strongly the can events ence dispersal relatively distance that reported long and infrequent rates, simulations spread mathematical potential Higgins explore used and to have (1998) Clark (1999) m. were Richardson 3 and mericarps than more that by found dispersed never 1989) (1984, Stamp simulations, certain of as distance such dispersal Geraniaceae, the that indicated (1999) Rooyen El 2005; 1980; al. 2006). al. et Albert al. et 1999; et al. Skorupa Mouden et Lane 1973; 1983; Soholt Ellner and 1930; Shmida Ridley rep- endozoochory; in or (myrmecochory), ectozoochory Hayward (by ants mammals by and birds transported tiles, or reported eaten, have lected, Others 1998). al. et that Burke R 1980; Noi-Meir 1976; and (Zeide capabilities dispersal nareetwt hi ifrn ye fncaisand nectaries of types different their with agreement In Diversifica- and Ecology, Pollination Evolution, Flower Van and Reede Van and 1989) (1984, Stamp However, Erodium hc r oehthge hntema (Magall mean the than higher somewhat are which , Erodium, , r iia oteaeaevle fohrangio- other of values average the to similar are Geranium, — Pelargonium Monsonia Pelargonium h iio-ln eainhpi nimportant an is relationship visitor-plant The and Monsonia Geranium and and Erodium ihsplcneldncaisaere- are nectaries sepal-concealed with sect. Monsonia rsnsacnieal iest in diversity considerable a presents Erodium Shr-ida ra,a elas well as area), (Saharo-Sindian eiap o ed)cnb col- be can seeds) (or mericarps Erodium Peristera slw ae nexperimental on Based low. is , Erodium Pelargonium nrhr eipeeand hemisphere (northern aemaual wind- measurable have and ö ih aedispersed have might Geranium c 97 98 Loria 1978; 1977, sch and Geranium Geranium ihde nec- deep with Erodium and . Pelargo- might Pelar- and ó n eo . .Mu F. C., Aedo, hog h eerhpoetRN0001/L n REN2003-04397/ and Spanish REN2000-0818/GLO the GLO. project by research financed the partly visits through our was during work assistance This kind Direcci loans. for specimen grateful herbaria also cited for are the and S. We of literature. curators Alpinar, and the K. specimens to L V. with and N. help and Guittonneau, for determination; G.G. Salamanca) Novoselova Gillespie, insect I. de with Egorova, with help (Universidad T. Castroviejo, for Portillo help Ciudad M. for Martin and Rodriguez A. Torres drawings; F. for figures; Castillo I. Gera- J. package; some analyses; Multidivtime with R8S the help with for help Navarro for Sanmart C. Roquet C. and sequences; Albers niaceae F. manuscript; the on eo . .J laoo n .Nvro 02 eiinof Revision 2002. Navarro. C. and Aldasoro, J. J. C., Aedo, httetnr shm omn oeDpeata large than Diptera more many to home is tundra showed the gradients latitude that and on altitude of Studies over formation found deterioration. the insects climatic the by and range, of ranges its of that mountain parts new (only in least high at triggered, is be genus Geraniaceae. this of of Pelargonium genera rate other diversification the The than latitude and altitude 1997). plants Menges and pollinators and between (Deyrup declines reduces reciprocal of usually risk generalization the of Moreover, limited sets 1983). were new specialization (Schemske these floral acquired of for possibilities Tertiary, many have the dominated of regions, must bees end opportunistic and As the times, pollinators. during more America or and once Europe of of clade The much 2002). al. and California, et Pons Fiz dium, 1991; 1981; Herrera from 1985; (Shmida resulted Quezel have regimes to likely disturbance is long-lasting which taxa, annual of portion higher a and pollination generalist to flexibility. shift of reproductive end a the by at Tertiary pollinators of the faced loss presumably and genera deterioration Those climatic the insects. of taxa that many flowers attract unspecialized produce ranges), American South 98 tpesne l 00 aize l 2004). al. et (Inoue Kalisz option 2000; evolutionary al. best et the Stephenson been 1988; outcrossing have and to of attraction seems as many pollinator rate In seen in Hemisphere. plasticity taxa Northern taxa, some the those of for parts reported in been colonizers in have rate changes autopollinaton Similar in increase who an pol- and (1988), inefficient linators of Inoue abundance pollinators between with relationship and agree a reported observations visitors These low-efficiency 3). of (Table spectrum shows rates) wide autogamy high a with them of many (and habitats 2000). al. bumble- of et Dlusskii and pollinators 2002; bees best (Kandori large the Thus, are 2000). bees effi- 1998; Ranker and Orthner and and activity (Bingham Bingham of temperatures low levels bees under high large ciency Totland only maintain 2002; However, bumble-bees 1999). Vokou and Olesen and and Elberling Blionis 1993; 3; (Table Hymenoptera A Geranium pro- high a by distinguished is flora Mediterranean The Erodium CKNOWLEDGMENTS 211 Geranium 205 47: sections ó – í eea eInvestigaci de General n o epwt IAaaye;M .Snesnfrhl with help for Sanderson J. M. analyses; DIVA with help for n 252. – h eu ihms pce rwn ndisturbed in growing species most with genus the , 297. zrlod,Neoandina Azorelloida, nldsmr pce dpe owd agsin ranges wide to adapted species more includes .(Geraniaceae). L. shge nGrnaee.Teseito could speciation The Geraniaceae). in higher is ñ zGreda n .Pno 98 ol hcls of checklist World 1998. Pando. F. and Garmendia, oz and h uhr hn .NeoFlnrfrcomments for Feliner Nieto G. thank authors The . Geranium L ITERATURE nlsdlJard del Anales and , ó Cient n ooie h Mediterranean, the colonised C Paramensia Geranium ITED í iayT y fica í Bot n (Geraniaceae). nteehabitats these in á iod Madrid de nico é nc (DGICYT) cnica Campanula ó e,adM. and pez, Geranium Blumea Ero- 56: . Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 08 I TA.PYOEY&BOEGAH FGRNAEE335 GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ of status generic The identification. 1996a. model F. statistical Albers, the at look new A 1974. H. Akaike, 2008] les .19b h aooi ttso h genus the of status taxonomic The 1996b. F. Albers, ae,B .P,A or n .A .Wlim.17.Gooyo the of Geology 1972. Helbr Williams. D. T., J. F. Bakker, A. L. and Mohr, A. P., H. B. Baker, S Ll. Vargas, P. Navarro, C. J., J. Aldasoro, mor- Anatomy, 2001. Aedo. C. and Vargas, P. Navarro, structur C. J., attracting J. 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Ge- yte 40: 40: 64: 15: 20: nd g s Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 3 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 336 es .E 98 otiuin oarvso of revision a to Contributions 1968. E. L. Kers, an importance pollinator various with visitors Diverse 2002. I. Kandori, ai,M,M hln .N aa,adR .Pnigo.20.Arc,the Africa, 2000. Pennington. T. R. and Labat, N. J. Seasonal Thulin, 1999. M. Naveso. M., A. Lavin, M. and Alonso, A. J. Alonso, C. J. J., S. Lane, hypothesis phylogenetic a and evidence for concern A 1989. G. diver- A. a Kluge, of Performance 2001. Bruno. J. W. and Thorne, L. J. H., Kishino, of rates evolutionary estimating for method simple A 1980. M. Kimura, ann,J .adP odlt.19.The 1996. Goldblatt. P. and C. J. the Manning, guises, many in comes Cupid 1995. Goldblatt. P. and C. J. Manning, Magall Phy- of Analysis MacClade: 1992. 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P. nearly W. Soltis, M. a S. and P. Wurdack, Soltis, J. D. K. D. Lled Sheahan, D. C. M. Johnson, M. A. S. Cameron, M. K. approach. likelihood Evolution penalized and a Biology times: divergence and trees. phylogentic Evolution reconstructing for method ,J,M srd,C Blanch C. Estrada, M. J., n, odcJunlo Botany of Journal Nordic ç – 9 259 59: Evolution 3in 43 – – e util i Munt del 164. 1574. :406 4: atpB ó ó eBarcelona de ete esp tres de sect. 5 257 55: 9 26 19: – ln osraini h eierna Area, Mediterranean the in Conservation Plant 264. and – 425. 4 1061 54: Jenkinsonia – – rbcL ’ Obac. 37. 309. 9 101 19: è salJra fBotany of Jornal Israel 9 33 29: eesequences. gene isend cies – salJra fBotany of Jornal Israel 1065. Science oorfe e evid ac auasde Naturals Parcs de Servei del Monografies – – é 0 in 109 (Geraniaceae). 109. n .Mlr.20.Booi ela de Biologia 2000. Molero. J. and , – 0 129 20: Ecology e eln ora fBotany of Journal Zealand New 43. 0 661 80: è rbc 1 874 31: 0:283 208: iusdlPr aua eSant de Natural Parc del miques Hypseocharis eunecomparisons. sequence L Coevolution eaimsanguineum Geranium iifrais(xod England) (Oxford, Bioinformatics – ln ytmtc n Evolution and Systematics Plant 140. – 1 1446 61: 671. ot fia ora fBio- of Journal African South – ó ytmtcBiology Systematic − eaimsessiliflorum. Geranium 879. – .C itu,M Powell, M. Pintaud, C. J. , onn ehd new A method: Joining 285. ot fia ora of Journal African South 0 105 30: oeua ilg and Biology Molecular – rbcL d .H Nitecki. H. M. ed. , Oaiaee iha with (Oxalidaceae) 2 147 32: 1459. eesequences. gene – 123. – 159. (Gerania- Molecular 9 306 49: 3 567 23: Annals ed.C. Mo- 1. – – - Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. etr .J .17.Amngahof monograph A 1979. T. of J. palaeoenvironment H. and Venter, origin The of 1975. distribution M. E. the Bakker, on Zinderen Observations Van 1969. M. E. Bakker, Zinderen Van del Venta de superieur Miocene du pollinique of Flore 1989. phylogeny E. The Campo, 2007. Van Albers. F. and Bakker, T. F. T., phenology, Touloumenidou, flowering Norway: alpine in Pollination 1993. O. Isles. Totland, British the of flora Biological 2004. J. R. rate Tofts, evolutionary and time Divergence 2002. of Kishino. H. rate and the L. J. Estimating Thorne, 1998. Painter. S. I. and Kishino, H. L., G. J. Thorne, D. and Jeanmougin, F. Plewniak, F. Gibson, J. T. D., J. Thompson, aelt . .Gel,G atu n .Bue.19.Uieslprimers Universal 1991. Bouvet. J. and Pautou, G. Gielly, L. P., parsimony Taberlet, using analysis Phylogenetic PAUP*: 2002. L. D. Swofford, etr .J .18.Pyoegah n neseisrltosisin relationships interspecies and Phytogeography 1983. T. J. H. Venter, 1999. 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R. g Smissen, El 2001. Grau. A. and C. A. Slanis, isfrmlil euneainetaddb ult nlsstools. analysis Research quality Acids by strate- aided Nucleic alignment flexible sequence interface: multiple windows for gies CLUSTAL_X The 1997. Higgins. h ai eetbiome. Desert Namib the Africa. in Ericaceae Moro. Evolution and Systematics Plant Monsonia. communities. plant Botany two of in Journal rates dian visitation and visitors, insect L. data. multilocus with estiation evolution. molecular of rate Evolution the of evolution ln oeua Biology Molecular non Plant three of amplification for Associ- Sinauer Sunderland: 4.0b10. ates. Version Meden- van methods), M. other J. (*and and Academic. Burgt Kluwer der Hague: van M. The X. Rooy. Maesen, de der bach van G. J. L. Monsonia Nederland Wageningen, Landbouwhoogeschool edlingen Plants ology. of species system pollination in reinforcement for shift. role a suggest data lutionary n.Jn:Gsa Fischer. Gustav Jena: ung. in considerations. ary gonium ytm n h reigsse of system self-incompatibility breeding (Campanulaceae). the the in and plasticity system, depresion, inbreeding among 78. Geraniaceae. 46. flora, pce of species Ecology of nal quences. .Svlie,W .Hh,S .Ho,M .Fy .Atl,S M. S. 2000. Farris. Axtell, S. rDNA, M. J. and 18S Fay, Nixon, from F. C. inferred K. M. phylogeny Kress, Hoot, J. Angiosperm W. B. Prince, S. M. L. Hahn, Swensen, H. W. Savolainen, V. Monographs ( rats Kangaroo of distribution bipolar the phyllaceae). for origin Pliocene Argentina. la Cali- County, Merced of areas fornia. agricultural in blackbirds tricolored ora fEcology of Journal Pelargonium Evolution eln Springer. Berlin: . caPalinologica Acta ot fia ora fBotany of Journal African South h Condor The Grnaee nSuhr fia clgcladevolution- and ecological Africa: Southern in (Geraniaceae) oaia ora fteLnenSociety Linnean the of Journal Botanical 5 1647 15: .(Geraniaceae). L. rdu.Junlo Ecology of Journal Erodium. Pelargonium 3 357 43: ü utainSseai Botany Systematic Australian ebooiceTpnasEeet e Sippenglieder- der Elemente als Typen tenbiologische 2 611 72: Darwiniana p 631 Pp. . 0 1596 60: iooy merriami Dipodomys – naso Botany of Annals 2 465 82: 1657. – ln ytmtc n Evolution and Systematics Plant ruet Geographica Argumenta 376. 2 537 92: – 1 1072 71: 620. 5 4876 25: ihntso t clg n olnto bi- pollination and ecology its on notes with – eiwo aaooayadPalynology and Palaeobotany of Review 7 1105 17: – :9 1: 3 in 638 1601. 9 343 39: – ora fBiogeography of Journal 467. – – Bothalia 32. 555. – − – 1079. h idvriyo fia Plants, African of Biodiversity The – ytmtcBiology Systematic 4882. oigrgoso hools DNA. chloroplast of regions coding é 1109. – nero 352. nteMjv desert. Mojave the in ) 5sp.:211 85(supl.): 4 865 14: Monsonia 7 1005 77: 6 467 56: rdu cicutarium Erodium Hypseocharis apnl rapunculoides Campanula 6 301 16: 6:1 264: ipra ilg fDesert of Biology Dispersal – 2 89 12: 869. – – 470. L.(Geraniaceae). oeua ilg and Biology Molecular 1020. eaimrobertianum Geranium – 3:381 133: – 14. Scleranthus – :65 2: 315. – 1 689 51: 219. rbc 97. 0:71 208: Oaiaee en (Oxalidaceae) 9 1 79: ,and L, – 73. – 461. – seeds. – 702. – 128. Ecological 97. atp (Caryo- 9 49 69: Cana- Pelar- Med- se- B Jour- eds. L. – mo Moulares, of Km os. UKY aay,2 mfo G from Km 27 Malatya, TURKEY, Boiss.: (MA-252363), oumo haapoic,bre fPtraprovince, Petarca of border (MO), province, Choapa Coquimbo, 4567 al. et Garmendia Yelmo, (Australia), El Desert Victoria DQ452878 DQ072044, Great in collected Coolgardie, DQ452877 and DQ452897, Roe between aminya, Mathez & Ficajola, DQ072051 UTAI,M.AnnBtncGarden, Botanic Annan Mt. AUSTRALIA, Esteban, cicutarium DQ452869 St Valley, Temescal Co., Riverside California, AJ235774. DQ452879 DQ452888 itmn .C 93 olnto in Pollination 1993. C. P. Zietsman, insect- in of patterns Dispersal syndromes 1976. B. pollination Zeide, ecological The 1986. T. Yumoto, 3 Pp Geraniaceae. of classification The 1990. F. P. Yeo, in type Fruit-discharge 1984. F. P. Yeo, organophylet ecology, structure, nectaries: Remarkable 1998. S. Vogel, (MA-393124), Aedo bold by indicated region. sequences (new num- for numbers her- herbarium type) accession voucher, and GenBank locality, in voucher and name, presented ber locality, taxon are Data sequence: their numbers. following including accession the GenBank study, and number present barium the in used e,P .17.Tebooyadsseaisof systematics and biology The 1973. F. using P. Yeo, inference phylogenetic Bayesian 1997. Rannala. B. and Z. usng analysis Yang, phylogenetic for package program in a display PAML: Floral 1997. 1979. Z. Rathcke. Yang, J. B. and Miller, J. L. F., M. Willson, the of antiquity the On 1983. Lancaster. Wikstr N. and Seely, K. M. D., J. Ward, PI,Mdi,Rzsd uroReal, Puerto de Rozas Madrid, SPAIN, Skhira, TUNISIA, Willd.: L Hamed, Bisegna, Pinnola, Rosa Mt. Abruzzo, atoij ta.14575 al. et Castroviejo Madura, of botrys W Km 59 way, DQ072065 oides DQ072029 é aionamacrophylla California A rdu absinthioides Erodium microphylla Balbisia aod Sessanio, de fano PPENDIX raniaceae). Botany meadow. alpine in plants pollinated Stellenbosch. of Symposium Univ. Geraniaceae International the Society implications. Linnean evolutionary the its and classification Society monifolia method. Carlo Monte Evolution Chain and Markov Biology a sequences: DNA likelihood. maximum and B) (serie London of tree. family the calibrating giosperms: Namib. cases. Miscellaneous prespectives. 90(MA), 6970 Ds. il:TNSA aKesra-Darsole, La TUNISIA, Willd: (Desf.) DQ072048 ö Cv)Bro. ... aiona a rnic,M.Tamalpais, Mt. Francisco, San California, U.S.A., Bertol.: (Cav.) ,N,V aoann n .W hs.20.Eouino An- of Evolution 2001. Chase. W. M. and Savolainen, V. N., m, er n ot4897 Bort and Serra G trnL eoe l 4931 al. et Aedo Geranium : ü , . ; , , , aar ta.2307 al. et Navarro OOC,Hg ta,Tiz-n-Test, Atlas, High MOROCCO, L)L (L.) ms1549 emes — .ciconium E. rsooabigelovii Crossosoma — — 5 221 25: 7 285 67: — .Smlsadacsin rvddfor provided accessions and Samples 1. ot fia ora fScience of Journal African South – ; Q586 DQ072034 DQ452876, ; nt,and Knuth, ; F ; DQ072073 .californicum C. .aureum E. .antariense E. .corsicum E. and ot fia ora fBotany of Journal African South ’ , H laoo3000 Aldasoro DQ459009 — dpaieaspects. adaptative : – – é 224. 346. .e io 1:SAN aaac,Fet eSan de Fuente Salamanca, SPAIN, (1): Aiton ex r. rbc ; (MA), .glaucophyllum E. Pi. ece HL,Atfgsa fPaposo, of W Antofagasta, CHILE, Reiche: (Phil.) (MA), eoe l 8108 al. et Aedo ; ; il. UKY us,Uludag, Bursa, TURKEY, Willd.: 6:2211 268: region. L .daucoides E. L)L (L.) 9 1 89: .cygnorum E. la. aar,Vra,S Vargas, Navarro, Aldas., (MA), (MA), , laoo3053 Aldasoro 4 717 14: aoi:ASRLA olade yeHig- Eyre Coolgardie, AUSTRALIA, Carolin: Ruberta optrApidBioScience Applied Computer — (MA-623612), DQ072035 L – rhr15 Archer Q708 DQ452881 DQ072078, , oy OOC,Hg ta,Tizi-n-A Atlas, High MOROCCO, Rouy: ’ ; 36. é H (MA-625205), Nutt. — a:FAC,Cria in,paedi plage Piana, Corsica, FRANCE, man: .crinitum E. é .Watson S. (MA), DQ072049 Q702 DQ452882 DQ072052, – r. . ‘‘—’’ – 2220. : 724. esm lucens Bersama Dum. os. PI,Je,Crio Nuevos, Cortijos Jaen, SPAIN, Boiss.: TL,Arzo L Abruzzo, ITALY, : ; eagnu dolomiticum Pelargonium es utvtdi Afo seeds from MA in cultivated Nees: rdu hirtum Erodium GenBank, .alpinum E. niae osqec o h DNA the for sequence no indicates DQ072016 L)L (L.) , (MEL-2039223), Geranium (MA), Evolution Flora rhr27069213 Archer — (MA), DQ072059 oaia ora fteLinnean the of Journal Botanical : clgclResearch Ecological ; , L S,Arizona, USA, rceig fteRylSociety Royal the of Proceedings Gillespie .geoides E. aoi:ASRLA Bur- AUSTRALIA, Carolin: ’ D d .Vse.Cp Town: Cape Voster. P. ed. , 9 175 79: ’ DQ072095 ó — Aquila, H ü 9:225 193: e 499 pez ’ uet405 Aubert Q709 DQ452875 DQ072039, r é ; 9 259 59: DQ072018 — Fern ü .e io:TNSA 14 TUNISIA, Aiton: ex r. Br . L f.) (Burm .brachycarpum E. oDarende, to n , Geranium Grnaee:isuein use its (Geraniaceae): 3 52 33: L11179. , csy,GenBank, Scyscyl, DQ452870 – ; laoo2842 Aldasoro á á 183. 0(MA), 10 , .arborescens E. Willd. dzCsse l 3277 al. et Casas ndez Timbal z&Ad:U.S.A., Aedo: & ez – — Conti .S.Hl:CHILE, St.-Hil.: A. – – oaia ora of Journal Botanical (MA), 248. ; , ’ 8in 18 264. – DQ072066 3 555 13: qia r Santo pr. Aquila, ; it eie 1580 Feliner Nieto .cicutarium E. — trnL 63. .cossonii E. (MEL-2015744), , (NSW-213829), sections , ; Á — ’ salJra of Jornal Israel H .gaillardotti E. alr10620 Taylor lvarez – 66(MA), 1656 rceigof Proceeding ..(MPU), s.n. ; :83 1: nt (Ge- Knuth ; F DQ072050, DQ072013, é .gruinum E. – . ITALY, r.: .aspleni- E. 556. Molecular and Mu – , (Desf.) Godr. SS137, 95. (MA), Guitt. — Phlox Ane- ñ rbcL ; ; (2): oz- — ï E. E. ic t- : , Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. rnh&Sv AA,Hnh,Tym-e,Ttym-h,M Tsurugi- Mt Tateyama-cho, dake, Toyama-ken, Honshu, JAPAN, Sav: Franch.& UKY Ermenek, TURKEY, al eAr de Valle C GenBank, rz,I Aquila, Il bruzi, dit, AF167151; (MA593417), 4144 DQ452902 AY944418 tum Sumelas, Kasdir, Trabzon, TURKEY, DQ452918 Nordm.: ex Ledeb. (1) Mts, Rhodope BULGARIA, B Atlantiques, 2433 Aedo DQ452886 L. Iblane, Bou Jbel Medio, catum etdi Chirrip in lected DQ452930 DQ452921 core-core — alsd aRapita, la de Carles DQ452931 Timhadit, de S Atlas, Middle ROCCO, etdi California, in Cape, lected the in californicum collected seeds from 4474 (Yemen), Attuba and Hilla between collected Falls, Rock DQ452925 Smooth USA, Britton: Erdek, Bandirma, KEY, 2018 al. et Herrero er,Itatiaia, neiro, DQ452885 DQ452926, CA3870 argenteum asnForresti Maison Kilimanjaro, Mt in L11184. utvtdi Afo ed olce nGeorgia, DQ452884 in collected DQ452915, seeds from MA in cultivated 7192 Aedo — — PI,C SPAIN, USA, Arizona, texanum Co., Yavapai in collected rmsescletdi ree,S Turkey, SE Ermenek, in collected seeds from stephanianum Er-Rachidia, of NW DQ072041 uprMkn) ewe aga n Yushu, and Jangkar DQ072027 between Mekong), (upper DQ072077 PI,Gaaaaa Ca Guadalajara, SPAIN, utvt nM rmsescletdi gp,CioSe eetRoad, Desert Cairo-Suez Egypt, in 3487 collected seeds Aldasoro from MA in cultivate io:SAN C SPAIN, Aiton: DQ452872 Letur, DQ452883 DQ072071, Igherm, Atlas, Nevsehir, dept. Vadisi, (MA), DQ452874 3 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC L (L.) 338 srveo15237 astroviejo rno appendiculata Francoa L14694. , ; ; : : E. TL,Arzo Ceppo, Abruzzo, ITALY, .cazorlense G. GenBank, eo4235, Aedo a. AER,ProMnz hoRibeira, Chao Moniz, Porto MADEIRA, Cav.: (MA), Est ’ UKY hmsae kDagh, Ak Ghumushane, TURKEY, : “ Á DQ072026 H variabile eaimaculeolatum Geranium vrz1239 lvarez (MA), eo4118 Aedo é .Ga 2:GenBank, (2): Gray A. é á . ODNA eas (Jerash), Gerassa JORDANIA, r.: ae 182 banez tu. HL,Atfgsa Toconao, Antofagasta, CHILE, Steud.: — , ; , , , , , , , ; ; , (MA), , i,Zhr el Sierra, la de Zahara diz, . utvtdi Afo ed olce nFrance, in collected seeds from MA in cultivated L.: (MA), .pyrenaicum G. á — — — — .jahandiezianum E. DQ452871 .guttatum E. — .dolomiticum G. — DQ452873 .L oe .L oe:cliae nM rmsescol- seeds from MA in cultivated Jones: L. F. & Jones L. G. — DQ452911 .lucidum G. ,Bagergue, n, AF167152; , ; ; il. HN,Qnhi Nangq Qinghai, CHINA, Willd.: ; ; é ; ; ; DQ072050 (MA), le 2762 Alves .cinereum G. .pelargoniflorum E. G — .columbinum G. .pusillum G. ” horl é .costaricense G. (MA593341), .biuncinatum G. .phaeum G. ó ó eAin-Almon re peue hybrid (presumed DQ452889 , L14696; , á laoo3443 Aldasoro i s.n. miz DQ452903 ot Rica, Costa , (MA643798), ewo:SAN an Cazorla Jaen, SPAIN, Heywood: i,Zhr el Sierra, la de Zahara diz, — (MA), ipr s.n. Aizpuru é (MA-591697), (MA593333), (MA690990), olc 43213 Podlech eo5450 Aedo guy, DQ452907, ; laooA9140 Aldasoro eao ta.9387 al. et Velayos DQ072023 ; .hoefftianum E. ; ; ; ; , .neuradiifolium E. Cav eo4199 Aedo .brasiliense G. atoij 15201 Castroviejo .texanum E. .atlanticum G. .sebaceum E. Ds. il. UII,Feriana, TUNISIA, Willd.: (Desf.) — . OOC,Xun blLka Hauta-el- Lakra, Jbel Xauen, MOROCCO, L.: G Mu , eo2966 Aedo eo4842 Aedo (BC), DQ452929 um . OOC,Mdl ta,STimha- S Atlas, Middle MOROCCO, F.: Burm. (MA), . ñ — Cav ; .() OOC,Tfet ta ei Aedo Medio Atlas Taffert, MOROCCO, (1): L. .: rotundifolium oh. PI,Le SPAIN, Rothm.: .nanum G. eo10351, Aedo , mrs Atienza, amares, .caffrum G. , ñ . RNE Xuberoa, FRANCE, L.: ; — HL,L Araucan La CHILE, Oliv zGredae l 4626 al. et oz-Garmendia DQ452908, — err ta.1995, al. et Herrero .austroapenninum G. m. ar ele:MRCO Anti- MOROCCO, Weiller: & Maire Emb., , . PI,Grn,Ser e Montsi del Sierra Gerona, SPAIN, L.: . tu. utvtdi Afo ed col- seeds from MA in cultivated Steud.: atoij 15137 Castroviejo DQ452927 (MA), DQ072022 ; uy15548 Jury : DQ452928 owr:cliae nM rmseeds from MA in cultivated Kokwaro: ; (MA), — , .sonchifolia F. os.&Hlr nBis:cliaei MA in cultivate Boiss.: in Heldr. & Boiss. PI,Hec,Ordesa, Huesca, SPAIN, DQ452901 DQ452913 — DQ452914 DQ072069 .endressii G. .: (MA593345), (MA-464889), — L14693; , aar 3425 Navarro utvtdi Afo ed collected seeds from MA in cultivated ; .Ga 1:cl.i Afo seeds from MA in cult. (1): Gray A. (MA593743), (MA), , .reichardii E. eo4472 Aedo , ; .A e. UKY G TURKEY, Mey.: A. C. eie OOC,Bui,8km 8 Boumia, MOROCCO, Delile: rgli at:BAI,Rod Ja- de Rio BRASIL, Mart.: in Progel .pusillum G. — DQ452905 DQ452919 .paularense E. os. OOC,Beni-Snassen, MOROCCO, Boiss.: eo4234 Aedo ose at:MRCO Atlas MOROCCO, Batt.: ex Coss DQ452909, cl eh:cliae nMA in cultivated Zeyh.: & Eckl. DQ072043 eiee or,SAN Albacete, SPAIN, Godr., ex Delile ; osna 43581 Cousineau (MA), . UKY vie,KzDagh, Kaz Evciler, TURKEY, L.: (MA626513), .asphodeloides G. laoo3492 Aldasoro — , , , (MA643260), DQ452923, — — , , ; — , , .tordylioides E. — — — ; ; aar 3424 Navarro .reflexum G. .Gy RNE Pyr FRANCE, Gay: J. — Wld)Cv:GenBank, Cav.: (Willd.) ; Vargas .cataractarum G. .malacoides E. laoo2838 Aldasoro eo4097 Aedo DQ452904 . ; ; ó AY944424 , ê .sylvaticum G. (MA), ; Q712 DQ452880 DQ072102, , x ryasutherlandii Greyia .oxyrhynchum E. , DQ452922 (MA685246), ,Pirna Ferradillo, Priaranza, n, in Wo Jangkar of NW Xian, n .brycei G. (MA593420), DQ452912 eo468 Aedo DQ452920 í eo TL,Abruzos, ITALY, Aedo: .macrorrhizum G. — ,8K fYupehue, of N Km 8 a, eo3864 Aedo DQ452887 , .corsicum), E. ags125PV98 Vargas .() GenBank, (2): L. oe l 2892 al. et Ho — — ; en oz Izco, & Gonz. Fern. G Valc ; od3126 Wood ; — (MA), eo7006 Aedo . .macrorrhizum G. DQ452917 eo2967 Aedo DQ452910 (MA), Aedo .robertianum G. DQ072026 DQ452924 albanum ; um . TUR- F.: Burm. , .tuberosum G. . TL,Ab- ITALY, L.: ags335pv02 Vargas , á .E Br. E. N. — Ds. L (Desf.) — cl370VV01 rcel (MA-588866), , , (MA643259), , laoo2973 Aldasoro (MA-685245), L)L (L.) , (MA593799), , — — ; — ; ; DQ452906 57(MA), 2587 ö DQ072037, 9150 os:MO- Coss.: DQ452916 DQ072100 DQ072033, . SPAIN, L.: .bicknellii G. .nodosum G. .yesoense G. ; ee Ask reme, ; ; GenBank, .palma- G. .divari- G. .gracile G. .Bieb.: M. .Bieb., M. ’ , , H , , (MA), (MA), .(1) L. (MA), Hook. — (MO), Aedo : — (MA), (MA), á — ’ — é é Aedo, H San , n .ex r. ; ; ; ; é é ; — — L., L.: G. G. G. es r.: L. E. E. , , , : , , , , eBn AF167142, GenBank L14703; GenBank, marifolia ania — L14704; Z95284, AF167146, GenBank, GenBank, Salisb., (Cav.) Z95280, GenBank, Willd.: O .vanderietiae M. adnBtnqeNtoa eBelgique, de National Botanique Jardin ciz AII,Nre,GetBcsaln clctrsn (W), s.n. Schlechter Clairmont, Buchsmanland Great DQ452894 Naroep, NAMIBIA, Schinz, eb OOC,Tojat ihAls fteM the of S s.n Atlas, Finckh High and Staudinger Taoujgalt, MOROCCO, Webb, Kirstensbosch — — bilobata Hypseocharis DQ452867 Transvaal, N AFRICA, SOUTH DQ452891, Richard: A. ex Meyer wkpud hncanyon, Khan Swakopmund, 421PV02 Oujda, ignorata ALGERIA, Boiss.: (Cav.) Kirstenbosch, Africa, South in collected seeds from BC in 9e4pv99 cultivate Harv.: & utvtdi Afo ed ae nSuhAfrica, South in taken seeds from DQ452893 MA in cultivated antariense: laoo949 Aldasoro (MA); alpinum alnifolium (MA); 3450 Kill al. et Navarro 121 Spitzberger Jabalc cedrorum Mt Granada, SPAIN, 582, X*, Real, Puerto de Rozas (MA); Madrid, SPAIN, 80, A*, 2935 al. et Aldasoro Bizerta, Castell ,27 OOC,Trudn,Hg Atlas, crassifolium High Taroundant, Ajaccio, MOROCCO, Corsica, X,1267, in collected plants MA, in Pinar, del Navas 474 Lazarides Salamanca, Spain, 136, Trevenque, Granada, SPAIN, 777, X*, Biespors, Stavropol, RUSSIA, 430, (D), X Dilar, Trevenque, Granada, (MA); 9193 al. et * 3 UTAI,OipcDm Mine, Damp rum Olimpic AUSTRALIA, 73, A*, Qarry, Axehead AUSTRALIA, Valley, cal herbarium from dioecy and P/O from deduced in sheets. was observation, dioecy autogamy and rest tests selfing the with automatic for Species gy- bagged unknown. (GD): and studied dioecious, index alloga- (D): P/O generally autogamous, n: generally X: nodioecious, A: types, facultative, F: Reproduction mous, Hypseocharitaceae. and ceae ’ A ; eagnu althaeoides Pelargonium ; AJ402972; , major Melianthus rdu absinthioides: Erodium aionamacrophylla California ign,Rancagua, Higgins, .rapaceum P. ysohrssp.: Hypseocharis í * 5 UTAI,Mukimbudin, AUSTRALIA, 75, A*, : PPENDIX ni, .corsicum E. .carolinianum E. ó .aguilellae E. arborescens: E H *() 9,IAY L ITALY, 890, (D), X* : , ,Od,Ser elsPedrizas, las de Sierra Onda, n, laoos.n Aldasoro .capitatum P. ex.&Schr. & Merxm. ö DQ072073 (MA), us:A,21 UII,Sejenane, TUNISIA, 221, A*, Guss.: cot&Ktcy D,34 UKY l a,Nidge, Dag, Ala TURKEY, 334, (D), X Kotschy: & Schott ad aae 7612 Hadacek & randl — , ; , (MA); * 1,MRCO Tizi-n-A MOROCCO, 519, X*, ilsi s.n. Gillespie Ds. DC. (Desf.) ods.n. Wood — — — .Ifre erdcintp n / nee nGerania- in indexes P/O and type reproduction Inferred 2. .multifida M. L14708. , (MA); (NSW); Cav. .major M. ; ; ; laoos.n. Aldasoro (W); ase 25-8-77 Gassner .camdeboense M. L o. les GenBank, Albers: Bol.) (L. .praemorsa M. L)L (L.) .emarginata M. DQ072010, laooadAlarc and Aldasoro -* uooptblt nrae natm,67 Cult. 617, autumn, in increases autocompatibility X-F*, : .acaule: E. , GenBank, 1:GenBank, (1): L L .chium E. (MA); .cicutarium: E. (MA); GenBank, , ó ’ ilp EU Cuzco, PERU, Killip: (W) .carvifolium E. (MA); — -* 4,EYT Avdat, EGYPT, 449, X-F*, ei. eBn,AF036065, GenBank, Herit.: e da,Fbea .Mateo G. & Fabregat Udias, pez — L)L (L.) laoo&a. * 4 UTAI,AesRock, Ayers AUSTRALIA, 94, A*, al.: & Aldasoro : 3:GenBank, (3): eo7065 Aedo * 2 ... aiona iesd o,Temes- Co., Riverside California, U.S.A., 52, A*, : ; laooadAlarc and Aldasoro * 3,TUNISIA, 231, F*, (MA). *() 3,TRE,UuDg,Bursa, Dagh, Ulu TURKEY, 736, (D), X* ; .whytei P. (STAUD), . E e.e nt)Albers Knuth) ex Mey. (E. DQ452892 — .radlkoferi G. .astragaloides E. : .chrysanthum E. L .pseudofumarioides P. DQ452900 ,73 TL,Sicilia, ITALY, 783, X, .asplenioides E. ; ’ L)Wld * 1,SAN od,Tavirana, Ronda, SPAIN, 316, A*, Willd: (L.) ’ aar ta.2246 al. et Navarro AII,Wnhe,Sossusvlei, Windhoek, NAMIBIA, ei. GenBank, Herit.: — .My xKuh OT FIA Natal, AFRICA, SOUTH Knuth, ex Mey. E. .cotyledonis P. ei. eBn,Z95299, GenBank, Herit.: — 2:GenBank, (2): ’ Mfe)Abr:cliae nSuhAfrica, South in cultivated Albers: (Moffet) Gardner-Torrens — qia Abruzzo, Aquila, (MA), ; L14707 , (W); .marlothii M. L14699. , * 9 PI,Salamanca, SPAIN, 59, A*, (MA685239), ags423PV03, Vargas Molero os.&Reut. & Boiss. akr eBn,AF167142, GenBank, Bakker, ó ó DQ072052, 8830 n , n, — .ciconium E. DQ452895 — — DQ452898 ; eoe l 7829 al. et Aedo AJ403027. , laooadAlarc and Aldasoro .emarginata M. laooadAlarc and Aldasoro ; ; zsy. GenBank, Szyszyl., ; Smabanova , od559 Dodd ile s.n Billier edi gracilis Wendtia .speciosa M. .exstipulatum P. os.&Reut. & Boiss. * 28 UII,Sejenane, TUNISIA, 1238, X*, : ó laoo3069 Aldasoro L)L (L.) BC-804188 AF167150 ï L 3470 n (MA); eo5256B Aedo Hamed, t laoos.n. Aldasoro ’ H amns.n Badman — laooe l 2865 al. et Aldasoro Eg. les NAMIBIA, Albers: (Engl.) é (MA); DQ459010 — , Knuth hri ta. 324 al., et Charpin (MA); — laooadAlarc and Aldasoro , . *() 9,GREECE, 397, (D), X* r.: ’ * 5 PI,Madrid, SPAIN, 95, A*, : ei. eBn,Z95294, GenBank, Herit.: , : L14701 Liston eoe l 8256 al. et Aedo DQ452868 ; .cazorlanum E. * 6,SAN Burgos, SPAIN, 761, X*, L14702; , — — (MA); (MA), . osnaangustifolia Monsonia (MEL); .major M. atoij ta.16583 al. et Castroviejo (LE); ; ; , , “ . utvtdi the in cultivated L., .beketowi E. : .redactum P. .nivea M. (MA), L14705 L i. L fil.) (L. : (MA); * G .aureum E. DQ452897, ” OT AFRICA, SOUTH eBn,Z95295, GenBank, (MA), : ete 2218 Penther (MA); ; HJ;TUNISIA, (HUJ); ü : (MA); . ó (MA); eecultivated, were .boissieri E. .brachycarpum E. Meyen * 4,SPAIN, 840, X*, ó mse l 1549 al. et emes .heliotropiodes M. * 9,SPAIN, 699, X*, — , Cv)L (Cav.) 9219 n .daucoides E. 9211 n DQ452896 — laoos.n., Aldasoro ; ; 2:GenBank, (2): .elongatum P. .botrys E. . .parvifolia M. laoos.n. Aldasoro ; DQ452899 — .australe P. ’ DQ452890 .crinitum: E. onMt., goun .gracilis, W. (Decaisne) ’ Heiwood ei.(1): Herit. .cossoni E. L11185. , ., * 102, A*, : Schmalh .cygno- E. (MA); (MA); (MA); — (MA); Aldasoro Vorster, CHILE, ó (W); ’ Vargas — Vargas 3477 n Herit. . Coss., Vivi- A*, : ; (W), , X*, : — M. E. E. E. E. E. E. : ; , , : : : : Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. edn rmsestkni ooc,Bn nse,Ad 41(MA); 5431 Aedo Snassen, Beni Morocco, austroapenninum in taken seeds from Reading Dagh Kav tarina, Ribadelago, Lake, L Bilheres, FRANCE, manca, ete, .Moe ,10 CAO,Ad ta.(2002); al. Creek, et Flat Alabama, Aedo USA, ECUADOR, 118, 170, A, Moore: E. 2288 (MA); MA, 1,SAN Ja SPAIN, 916, caeruleatum Attuba, MA, in Cult. P 140, Magallanes, CHILE, 96, A, azorelloides 5888 Gillett and Stauffer 2762 (MA); (MA); s.n. Shaw Airy 4379 Rohnia, TUNISIA, man-Ermenek, 13801 Jury 5022 Formentor, Mallorca, Spain, in collected Alarc plants MA in Cult. (MA); Boiss. mouretii L (L.) macrocalyx (MA); procedence unknown Lapeyr. MOROCCO, MA, in Cult. 258, A*, Range, Fraser AUSTRALIA, (BC); L Palomas, las de Puerto losum 339 Titeki, Agadir, GERANIACEAE OF BIOGEOGRAPHY & PHYLOGENY AL. ET FIZ Moulares, TUNISIA, 173, Valdecebollas, cia, glandulosum Lora, de Sargentes Burgos, SPAIN, 563, 2008] tibetanum s.n. Dniepopetrovskaia, christi UKRAINE, 498, (D), Alarc and Aldasoro Oujda, chum MOROCCO, 102, A*, hir, seeds from garden MA in 110), Cult. (sd: 9.2), 294 Azrou, (sd Morocco, april: 87 in 106); october: taken (sd: 63.7), 208 (sd: mean: 144 P:O june: autumn, in increases ibility Karaburun, KEY, Gerassa, DANIA, 4567 al. Alarc foetidum E. oe al. et Ho stephanianum: (MA); (MA); pelargoniflorum Ca Guadalajara, SPAIN, 1023, ane, by collected Kabili, drickii Tunisia, in taken plants ’ é H eaimaculeolatum: Geranium ia rm irad Gurp, de Sierra Trem, rida, é (LE); laooadAlarc and Aldasoro Mu lia s.n. Alpinar .e io:A 8 PI,Formentera, SPAIN, 68, A, Aiton: ex r. ó ó * 6 REI,Aaa Mts., Aragat ARMENIA, 76, A*, : eo4475 Aedo (MA); (MA); (MA); .tordylioides: E. (MA); : ’ L 9222 n 9213 n rs le 6377 Olsen Frost Sennen lia:X() 3,TRE,Lazistan, TURKEY, 433, (D), X Alpinar: H .reichardii E. .arachnoideum G. .albicans G. .recoderi E. .sebaceum: E. .caespitosum, G. eer as 6170 Pabst & Pereira ,91 PI,Almer SPAIN, 991, X, ’ : ñ H laooadAlarc and Aldasoro (MA); (MA); é Pit od3126 Wood * 8,SAN L SPAIN, 684, X*, ze l 4626 al. et oz .e io:A 8,SAN ard laoosn (MA); s.n. Aldasoro Madrid, SPAIN, 181, A, Aiton: ex r. .texanum E. é de.i ok f. Hook. in Edgew. -,14 COLOMBIA, 184, F-A, : G L (G. (SEV); . ,54 TL,Puglia, ITALY, 574, X, r.: .: atoij 15220 Castroviejo L)L (L.) .caffrum G. .touchyanum E. Schur .aequale G. Cv)Willd. (Cav.) oterts.n. Montserrat (MA); .arboreum G. (MA); *15,SAN ula lne oeo9(MA); 9 Moreno Alange, Huelva, SPAIN, X*,1255, : * 9,Cl.i Afo ed ae nQnhi China, Qinghai, in taken seeds from MA in Cult. 195, A*, * 0,TRE,Ermenek-Anamur, TURKEY, 603, X*, : (MA); é .tataricum E. * 6,SAN Castell SPAIN, 464, X*, .geoides E. laooe l 9119 al. et Aldasoro (MA); Blanch ó ,Nv eSPedro, S de Nava n, * 7,IAY Abruzzo, ITALY, 976, X*, : (ISTE); e)L pez) ’ uiut&Git1 * 78 PI,C SPAIN, 1728, X*, Guitt.1: & Auriault : H .trichomaniifolium E. .S-i. -,10 rga,Timote, Uruguay, 160, A-F, St-Hil., A. laooe l 9178 al. et Aldasoro agss.n Vargas ó eos.n. Aedo Mra)DC. (Murray) *(D,59 ugra tVitosha, Mt Bulgaria, 529, (GD), X* laooe al. et Aldasoro laooadAlarc and Aldasoro .gaillardotii E. .rupestre E. 5015 n é * 4,Mrco Aguelmane, Morocco, 846, X*, * 14 PI,Cdz Zahara, Cadiz, SPAIN, 1124, X*, (MA); * 4 S,Txs Post, Texas, USA, 74, F*, : laooadAlarc and Aldasoro r. .albanum: G. ,22 ut nMA, in Cult. 242, X, : é : .brycey G. ó Bb)Ad:A 4,GETBIAN Circester, BRITAIN, GREAT 146, A, Aedo: (Bab.) -,nUA ooao la Creek, Clear Colorado, n,USA, X-F, (MA); laooadAlarc and Aldasoro .St-Hil. A. ó 9216 n * 3,SAN eul Javalambre, Teruel, SPAIN, 737, X*, ta.9376 al. et (P); (MA); .hoeftianum E. e da ta. * 1,SAN una Tragac- Cuenca, SPAIN, 414, X*, al.: et Udias pez ,3,CIE Coquimbo, CHILE, 32, A, : -* 4,TNAI,Kilimanjaro, TANZANIA, 440, X-F*, : é .Gray A. laoo3043 Aldasoro uys.n. Jury * 1 (Sim 516 X*, eiee Godr. ex Delile Willd .bohemicum G. (MA); ó ia Senet, rida, .aristatum G. (MA); (MA); (MA); . (MA); 3472 n em s.n. Helms : (MA); ° ñ í .asphodeloides G. ,10 HN,Laddakh, CHINA, 160, A, Pur xCv)Cadevall Cav.) ex (Pourr. ,ClrAt,Adsr n Alarc and Aldasoro Alto, Calar a, aesadVald and Mateos (MA); eo4782 Aedo .jahandiezanum: E. Natales, amares, * 5,Cl.i MA; in Cult. 456, X*, : *() 5,TRE,Malatya, TURKEY, 353, (D), X* : imn 13019 Diamond : : .: 03(MA). 5033 -* uoetlt ihri tm,540, atumn, in higher autofertility X-F*, * 8,GOGA Kiziki, GEORGIA, 880, X*, -,n RSL Itatiaia, BRASIL, n, X-F, .recoderi E. (MA); (SEV); ,90 USA aai,Abakan, Jakasia, RUSSIA, 980, X, .brasiliense G. .brevicaule G. .biuncinatum G. : eoe l 20;G berteroanum G. (2002; al. et Aedo M) utvtdi Agre from garden MA in Cultivated (MA); .guicciardii E. tibr s.n. Steinberg .atlanticum G. (MA); ,80 AA,Mu,Haleakala, Maui, HAWAI, 840, X, * 0,TRE,G TURKEY, 108, A*, : .malacoides E. oin 3191 Soriano L ó (MA); ó 8832 n err s.n. Herrero ’ ó laooadAlarc and Aldasoro (MEL); laooadAlarc and Aldasoro ó ,84 AEOI,Mavrovo MACEDONIA, 804, X, : H ,Pe n, eo7468 Aedo ta.20) 9,SAN Palen- SPAIN, 690, 2000); al. et n 8837 n (MA); .trifolium E. laooadAlarc and Aldasoro ó (MA); é .janszii E. . -* 1,SAN Zamora, SPAIN, 216, A-F*, L.: 5019 n : . *() 1,TRE,Kara- TURKEY, 512, (D), X* r.: eyoas.n. Deryiova .manescavii E. -* 7,MRCO Taza, MOROCCO, 371, A-F*, eo4472 Aedo .guttatum E. S ñí err ta.2073 al. et Herrero *14,TRE,Evciler, TURKEY, X*,1048, : 2 Aldasoro á (MA); : scola, .laciniatum E. é zs.n. ez -,30 RSL .Ca- S. BRASIL, 360, X-F, : Aucher-Eloy .gruinum E. ok:AF,16 ut in Cult. 136, A-F*, Hook.: (MA); Garc s.n. s * 1,SAN M SPAIN, 712, X*, .ruthenicum E. (MA); os. *() 1,TUR- 312, (D), X* Boiss.: (MA); (MAF); -* 0,MOROCCO, 506, X-F*, ,3,YMN Hilla- YEMEN, 32, A, : * 7,SAN Sala- SPAIN, 175, A*, : Boiss (MA); (MA); (MA); .carolinianum G. Alarc í Cv)Cv:X,1335, X*, Cav.: (Cav.) eeas.n. Devesa Mart & a .glaucophyllum: E. laooadAlarc and Aldasoro laooadAlarc and Aldasoro alrs.n. Taylor (MA); (SEV); eo4474 Aedo .cataractarum: G. 04(MA); 3024 laooe l 9150 al. et Aldasoro (MA); .maritimum E. .: : .rupicola E. * autocompat- F*, : Alarc * 09 SPAIN, 1049, X*, (LE); ó .neuradiifolium E. aglfMrs.n. Mir Margalef .paularense: E. ó ó .bicknellii G. .cinereum G. ,70 ut in Cult. 710, X, osnut5290 Rossengutt 8838 n netal. 8835 n á os:X,722, X*, Coss.: * 9,JOR- 198, A*, : ö le Mar & Alves ó i,Tavirana, diz, P;Gumush- (P); Cv)Willd.: (Cav.) ee Nevse- reme, 8831 n .moschatum E. it Feliner Nieto laooand Aldasoro laooand Aldasoro .oxyrrhyn- E. .campii G. ó ahkas.n. Sachokia í .Be. X Bieb.: M. ta.203 al. et n .sanguis- E. n .lucidum E. (SEV); (MA); (MA) Mu .nervu- E. (MO); (MA); Schlieben (MA): (MA); : ó (Boiss.) * 44, A*, .hen- E. 9218 n . A*, L.: (MA); á Colla: ñ Titov laga, A*, : X*, : zet oz (L.) ;G. X*, X*, F*, H. ó ó G. G. E. E. E. E. E. E. í n n n : n:XF 5,SNWC ISLANDS, SANDWICH 256, X-F, ing: s.n. Pabot PER Chirrip Rica, (MA); Alarc columbinum: G. Castrovalnera, Santander, SPAIN, 1020, otert922 Montserrat Guaranda, Chimborazo, dressii: ECUADOR, 230, F, Valdelosa, de Villanueva Leon, Ghumushane, Valderrobles, PER 628, X, collinum G. um . * 5,SUHARC,Cp,KrtnohGarden, Kirstenboch Cape, AFRICA, SOUTH 452, X*, f.: Ermani-don, Burm. Ossetia, S GEORGIA, 1096, X, paz, Cipsira, losericeum Dzyra-Mts. Mts. Abjasia, GEORGIA, Trabzon, TURKEY, 669, Engler 2401 fallax G. s.n G) ,KZKSA,AaTau, Ala KAZAKHSTAN, n, (GD), X LEBANON, 846, X, Schenk.: Keibrouan, Feinour, LEBANON, libani G. nosum Kosh-e-Yelag, Iran, in collected seeds from MA (P); in collector Cult. unknown Kilimanjaro, NIA, Trujillo, 9097 al. ocnAtsn,L Mica, La Antisana, Volcan Gecidi, josan Hilario, J. Pico (MA); eo7224 Aedo magellanicum G. maderense G. (P); 46 Boissier Lydia, KEY, Coru SPAIN, 69, Korea; S Kaybang, in Compton krameri G. ard Montejo, Madrid, MA PER 152, A, Kost, CHEKIA, 816, X*, Nevada, 1143 bault 5202 Olsen Frost al. et Aedo 6,Cl.i Afo ed olce nRsi yKrutscherovskaya, (1991); by Herrera Russia Rize, in 860,TURKEY, (GD), collected X* seeds from Irkusk; river, MA Turyga in Cult. 568, Rize, s.n soro Jajce, BOSNIA, Mesolongiou, GREECE, ECUADOR, A,166, Benth.: Oman, in collected al. et Aedo paludosum matum Zeyh. Ta-tien-lou, Tibet, CHINA, 620, USA, in collected seeds 1955 al. et rero Shireteki, PER Guirigai, Trujillo, VENEZUELA, Moupin, 378, Tibet, CHINA, 686, X, PI,Salamanca SPAIN, (MA) . Ú Ú ó Alarc erz ta.320 al. et Pedraza (MA); iae l 893 al. et Nisa : , 8627 n , * 0,PRUA,Mdia Cantias, Madeira, PORTUGAL, 700, X*, : : .foreroi G. .costarricense G. a. -* 9,SAN acln,Matar Barcelona, SPAIN, 196, A-F*, Lam.: eoe al. et Aedo ,40 OT FIA Queenstown, AFRICA, SOUTH 460, X, * 06 ut nJc rmsescletdi .Prno,France, Pirineos, E. in collected seeds from Jaca in Cult. 1086, X*, ,30 RSL a al,Jordao, Paulo, Sao BRASIL, 310, F, Aedo (MA); tu. -,14 OII,Potosi, BOLIVIA, 124, A-F, Steud., (MA); in tal. et Riina .H ai:X 50 UKY Iskenderun, TURKEY, 1560, X, Davis: H. P. .exallum G. ; Smelsson (G); Est ó (MA); il.e Spreng ex Willd. (MA); .Kuh ,20 COLOMBIA, 280, F, Knuth, R. rnh a. * 1,Cl.i Afo ed olce by collected seeds from MA in Cult. 618, X*, Sav.: & Franch. (2002); n laoos.n Aldasoro and n (2002); tpa xWilld. ex Stephan .favosum G. Ú Ú ó é e:X,52 AER S,Bc eCana, de Bica IS., MADEIRA 562, X*, Yeo: (MA); (2002); , eoe l 5077 al. et Aedo ae s.n. banez (MA); eoe l (2002); al. et Aedo , laooe l 9168 al. et Aldasoro .cuneatum G. , .platypetalum G. err ta.1467 al. et Herrero .jahnii G. (MA); oeo635 Forero eoe al. et Aedo * ,SAN Castell SPAIN, n, A*, rc 3075 Krach eo ,F 4,CLMI,S eaad at Marta, Santa de Nevada S. COLOMBIA, 442, F, X, Aedo: .pyrenaicum G. ok . -,14 HL,L ruai,Volc Araucania, La CHILE, 144, A-F, f.: Hook. eoe al. et Aedo M)adcl.i MA; in cult. and (MA) .malviflorum G. ñ (2002); .makmelicum G. laoos.n Aldasoro a, .macrorrhizum G. (G); .mascatense G. Alarc 9 (MA); 597 eo6083 Aedo .core-core: G. (MA); mg s.n Amigo .oreganum G. .E or:FA 1,ECUADOR, 218, F-A, Moore: E. H. Alarc .nepalense G. -* 2,Cl.i Afo ed ae nCosta in taken seeds from MA in Cult. 428, X-F*, : .erianthum G. .maniculatum G. tu. ,30 EAO,Sinpe, LEBANON, 340, F, Steud.: (MA); (MA); atoij ta.15640 al. et Castroviejo tnl:F 6,VNZEA P VENEZUELA, 266, F, Standl.: eos.n. Aedo .crenophillum G. acre 370VV01 Valcarcel ó n laoos.n Aldasoro and n (MA); ó .potentilloides G. ile 11132 Nielsen (2002); ags&Narv & Vargas subsp. n laoos.n Aldasoro and n M) ugra tVitosha Mt Bulgaria, (MA); eoe l ( al. et Aedo .: .maculatum G. (2002); Pabot iae l 765 al. et Nisa (MA); ,48 OOBA udnmra Suma- Cundinamarca, COLOMBIA, 418, X, : (MA); . -* 6,SAN Santander, SPAIN, 562, F-X*, : ic.e .A e. * 31 TURKEY, 1371, X*, Mey.: A. C. ex Fisch. , (MA); . .nodosum: G. (MA); .humboldtii G. * 3,SAN una U Cuenca, SPAIN, 736, X*, .kilimandscharicum G. os.&Ru. ,68 PI,Gaaa S Granada, SPAIN, 668, X, Reut.: & Boiss. (MA); Vald -,19 HL,Antofagasta, CHILE, 159, A-F, Alarc ai 1870 David eo ,76 EAO,Cedres, LEBANON, 756, X, Aedo: rk 553 Drake .glaberrimum G. os,F,28 ut nM rmseeds from MA in Cult. 288, F*, Boiss, Howell (MA); (MA); obut1145 Gombault .dissectum G. (G); hololeucum * 7,COTA Krasno-Jablanac, CROATIA, 472, X*, : we:FX 2,JPN Hokkaido, JAPAN, 228, F-X, Sweet: .crassipes G. C:X 2,RSI,KrlsIs., Kuriles RUSSIA, 822, X, DC.: (MA); .divaricatum G. é .peloponnesiacum G. ó 9792 s .pseudosibiricum G. .molle G. eihvs.n. Levichev ó ey628 Remy ,Pe n, .macbridei G. n laoos.n. Aldasoro and n 2002); .pratense G. .linearilobum G. Dorr .E Moore E. H. os. ,94 EAO,Ehden, LEBANON, 994, X, Boiss.: .macrostylum G. .orientali-tibeticum G. : .dolomiticum G. laooadAlarc and Aldasoro . *(D,16,Cue (1977); Cruden 1062, (GD), X* L.: á *(D,12,Cl nM from MA in Cult 1326, (GD), X* eoe al. et Aedo (MA); (UPA); L z2151 ez eer 337 Texeira .glanduligerum G. cut 92 Schulte * 0,IAY Abruzzo, ITALY, 702, X*, ñ (MA); (MA); (MA); ohn s.n. Gochina ’ .kotschy G. (MA); agolosa, (MA); (MA); . e xDC ex Her 13(MA); 9133 Spreng .nivale G. A ry alus Biss- & Carlquist Gray) (A. (MA); L.Geranium:molle uen11074 Luteyn oaosys.n Kolakovsky ok xA ry ,208, F, Gray: A. Ex Hook. . ,12 pi,Teruel, Spain, 112, A, L.: (P); * 0,TRE,Kara- TURKEY, 902, X*, : opr434, Cooper .purpureum G. (MA); ó (MA); .ornithopodon G. (MA); .gymnocaulon G. Ehrh. .ecuadoriense G. eia2448 Medina Aedo .moupinense G. , .phaeum G. L.. .multiceps G. .digitatum G. n. ,9,TANZA- 90, A, Eng.: : : .psilostemon G. oterts.n Montserrat .pavonianum G. (2002); M) Calacoto, (MA); eo&a.s . sn al. & Aedo ,16 ECUADOR, 176, A, ,20 ECUADOR, 240, F, yegr45264 Nydegger : C nLm DC.: & Lam. in DC. (MA); .Knuth R. os,X G) n, (GD), X* Boiss, * 10 SPAIN, 1100, X*, .: .Myr *(GD), X* Mayer: J. ,17,SPAIN, 1078, X, : : os. ,n TUR- n, X, Boiss.: (MA); Alarc : .lucidum G. .libanoticum G. eoe al. et Aedo á .ibericum G. ,2,TURKEY, A,124, eo9696 Aedo Boiss eo9888 Aedo .multipartitum G. (MA); * 1,Cl.in Cult. 112, A*, -,36 PER 396, X-F, ñ aod Jab de ramo oeae 1854 Hohenaker a, ó .palustre G. (MA); 9201 n (MA); .gracile G. (LE); . .Kuhin Knuth R. .Knuth R. ó laooand Aldasoro Alarc * 1020, X*, : n Alda- and n il:A 61, A, Vill.: .incanum G. á laooet Aldasoro : : eo7073 Aedo uc. X, Turcz.: .lanugi- G. .Knuth: R. Llaima, n : (MA); -,416, X-F, ,1185, X, Hieron.: Franch.: ,946, X, : * 152, A*, cl & Eckl. (MA); . Ledeb. (2002); .pal- G. ó Bricq . A*, L.: .en- G. (MA); (MA); (MA); (MA); .ho- G. and n Gom- Cav.: Gage Her- Beck X*, : (G); : ó L., Ú A. G. X, n, .: ª , : Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ro 2220 Krook brevirostrata attenuata M. Jumbaya, Nercha, river Zabalkaskaia, Siberia, RUSSIA, 862, ed olce nMrco augl,At Atlas, Alto Taoujgalt, Morocco, in collected seeds (COI); 4734 Slechter Berg, Uporne Natal, AFRICA, SOUTH Violdrof, (W); Lasgtu, folia Great BOTSWANA, 1007, X, Knuth: wayo, (MA). Park, Kagasuyama Komado, Toyama, JAPAN, Gallatin. Mt Montana, (MA); (MA); tuberosum: tovarii laoo94,9053 9049, Aldasoro salmoniflora Aus-Witputz, AFRICA, SOUTH 4965, (MA); X*, Seeds Silverhill by provided seeds from parvifolia AOVRE Maio, VERDE, CABO praemorsa (MA); Garden Kirstenbosch (COI); Albers Oliv.) Karas, Grunau-Klein 1012 Paulo and Garies-Bitterfontein, AFRICA, SOUTH 9078 5730, X*, Albers: (Sweet) Garden, Kirstenboch seeds, Silverhill by supplied seeds from den London, East Africa, AFRICA, SOUTH Steinkopf-Violdrof, AFRICA, seeds, 491 Cooper Mu & Aedo Link.. ex Tchen-keoutien, India, CHINA, in 148, collected A, seeds from P in Cult. 446, X*, Sweet: Paxton & Lindl. ex Siebold Yunnan, in taken seeds from Garden Wakehurst China, in ECUADOR, Cult. 232, 142, F-A, Knuth: A, Planch.: & Triana Carolin Park, Kagasuyama Hiroshima, JAPAN, 7486 Aedo (2002); nt nEg:X,44 uti Afo ed olce nCia Yun- China, in collected seeds from MA in Tsen, Kia Cult Ou 494, nan, X*, Eng.: in Knuth 5156 santanderiense wkpud hncanyon, Khan Swakopmund, weddellii G. Alarc et suzukii Aldasoro G. Lunada, la de Portillo Santander, SPAIN, Campoo, Papudo, Valparaiso, CHILE, Torul-Trabzon, sibiricum sanguineum rdn(1977); Cruden robertianum: Trautv. Moore E. Villanos, Pico Tenerife, SPAIN, Waimea, Ossetia, N 4 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC 4219 al. et Navarro s.n. Aldasoro 340 osnaangustifolia Monsonia ysohrspimpinellifolia Hypseocharis nt:X 85 OT FIA aa,Alexandra, Natal, AFRICA, SOUTH 3835, X, Knuth: (MA); (MA); .herrei M. en 23 Heany eo ,F 5,Per 254, F, A, Aedo, .viscossisimum G. .versicolor G. Aldasoro eo3886 Aedo .multifida: M. .sessiliflorum G. : : ,n AII,Naroep, NAMIBIA, n, X, : : G) 6,USA,Clrd,DlaCo, Delta Colorado, U.S.A., 864, (GD), X ,10 UTAI,Hume, AUSTRALIA, 150, A, : ,30,SUHARC,Ntl Clairmont, Natal, AFRICA, SOUTH 3900, X, : . ,n AA,Hokkaido, JAPAN, n, X, L., oaa3590 Hosaka laoo9042 Aldasoro laooadAlarc and Aldasoro ,n USA .Baikal, L. RUSSIA, n, X, etr 4952 Venturi (MA); ,14 COLOMBIA, 164, A, Masam *(D,n UKY ruu,Plnoe a,Hreo1732 Herrero dag, Palandoken Erzurum, TURKEY, n, (GD), X* (W); CI;cl.i Agre rmsessple ySilverhill by supplied seeds from garden MA in cult. (COI); Bricq .sericeum G. uc s.n Busch .longipes M. L. ñ Mfe)Abr:X,40,SUHARC,Aus-Witputz, AFRICA, SOUTH 4601, X*, Albers: (Moffet) av:X 65 OT FIA Natal, AFRICA, SOUTH 1605, X, Harv.: nt:AF,28 OT FIA arsih Tafelroop, Harrismith, AFRICA, SOUTH 288, A-F*, Knuth: : * 1,SAN Santander, SPAIN, 110, A*, oz-Garm. (MA); : * ,AGL,Mossamedes, ANGOLA, n, X*, .Kuh ,64 OOBA Cundinamarca COLOMBIA, 624, X, Knuth: R. -* 1,SAN aar,Petilla, Navarra, SPAIN, 612, F-X*, B ; (B) L o. les ,45,SUHARC,Steinkopf- AFRICA, SOUTH 4150, X, Albers: Bol.) (L. (COI); .ruizii G. iae l 712 al. et Nisa ..(MA); s.n. .burkeana M. .: .sibbaldioides G. (MA); .: laoos.n. Aldasoro ,50 BOLIVIA, 560, X, (MA); atoij 14931 Castroviejo L. (MA); .luederitziana M. * 1,Cl.i MA, in Cult. 212, F*, .reflexum G. * ,cl.i AGre rmsescletdin collected seeds from Garden MA in cult. n, X*, (LE); . hlisn379 Phillipson : laoo9074 Aldasoro laoos.n.(MA) Aldasoro a. * 4,CIE aalns uroArenas, Puerto Magallanes, CHILE, 140, A*, Cav.: .camdeboense: M. ads eMts6269 Matos de Cardoso : * ,SUHARC,Albert, AFRICA, SOUTH n, A*, : * 5,IAY iii,Nebrali, Sicilia, ITALY, 954, X*, il.e Spreng. ex Willd. (P); (MA). ic.&C .My nC .Mey. A. C. in Mey. A. C. & Fisch. nt:X 6,KNA Nairobi-Namanga, KENIA, 364, X, Knuth: umr,9312 Summers, .subargenteum G. ,14 OTGL aer,Cnis A Cantias, Madeira, PORTUGAL, 194, F, (MA); .renardii G. Hieron .senegalensis M. ú .reuteri G. .ciliata M. : , .retrorsum G. lnh xHr. ,53 IBBE Bula- ZIMBABWE, 563, X, Harv.: ex Planch. ó .Remy J. eo6824 Aedo .nivea: M. eoe al. et Aedo -,10 ut nMA, in Cult. 180, F-A, 9196 n laoo9038 Aldasoro Vargas (MA); .inerme M. * 6,IAY ecr,Vd iSole, di Vado Pescara, ITALY, 766, X*, : M); (MA) aar ta.3098 al. et Navarro Benth eoe al. et Aedo .: agss.n. Farges -,22 PER 232, F-A, clctrs.n. Schlechter Komovalov US;SuhArc,cl.i AGar- MA in cult. Africa, South (UPS); ruv nTat. * 40 RUSSIA, 1430, X*, Trautv.: in Trautv. ok ciz * 07 NAMIBIA, 1067, X*, Schinz: & Focke (MA); (MA); eoe al. et Aedo : .: (MA); .sintenisii G. 08 RETN,Jji Volc Jujui, ARGENTINA, 6048, X Mfe)Abr:X,73,SOUTH 7337, X*, Albers: (Moffet) eo&Mu & Aedo (MA); * ,SUHARC,Craddock, AFRICA, SOUTH n, X*, * 2,cl.i Agre from garden MA in cult. 128, A*, ,10 PER 150, A, (MA); (MA); * 44 PI,Snadr Alto Santander, SPAIN, 1464, X*, : .skottsbergii G. (2002); ann 4410 Canning eoe al. et Aedo ; : Estebanez Est ,F 5,ECUADOR, 356, F, X, L .lanuginosa M. Rh. les * ,Cl.in Cult. n, X*, Albers: (Rehm.) ulei erte:A,193, A*, Perrottet: & Guillemin eo4779 Aedo .emarginata M. (2002): ’ .marlothii M. e xD. ,8,HAWAI, 80, A, DC.: ex Her laoos.n. Aldasoro é (MA); Aldasoro .sylvaticum: G. ae s.n. banez mt 3258 Smith .mossamedense M. (MA); (MA); .soboliferum G. .wallichianum G. xdladMendon and Excdel (MA); .pattersonii M. (2002); Est .transbaicalicum G. laoo9054 Aldasoro Ú W;cl.i AGarden MA in cult. (W); ilvnk 1755 Siplivinski ry:X 9,TURKEY, 996, X, Freyn: tuigrs.n. Staudinger (MA); é uahv1153 Sukachev .richardsonii G. , ñ ipr 3865 Aizpuru (MA); ae s.n banez Ú .wlassovianum G. (2002); zGr. -* 362, F-A*, oz-Garm.: .speciosa M. eoe al. et Aedo .crassicaule M. , ..(MA).; s.n. oprs.n. Cooper .yesoense G. eo3046 Aedo (MA); eoe al. et Aedo .wilfordii G. .Kuh * 164, A*, Knuth: R. (L); (MA); ,n NAMIBIA, n, X, : ods.n. Wood err 1431 Gerrard Garc .rhomboidale G. (MO); * 80 South 2840, X*, : eo5170 Aedo nt:X 3245, X, Knuth: M) 140-230, (MA); .sinenense G. : .strictipes G. .stramineum G. ó (MA); . *(D,820, (GD), X* * 9,USA, 994, X*, o. ,138, A, Kom.: D. Albers: (DC.) 29(MA); 9209 n .thunbergii G. uai 1342 Rudatis í ta.1481 al. et a .l M. .solanderi G. .Dnex Don D. Wl.ex (Welw. ,11753, X, : (MA); .grandi- M. (2002); eoe al. et Aedo Fern (2002); ,1008, X, : (MA) (MA); .glauca M. (MA); (MA); (MA); (MA); ç Serg ic.& Fisch. Maxim d 6694 edo Aldasoro ´ (W); * n, X*, : 21775 a heritieri .yeoi G. Polhill (MA); Fisch. á (W) ndez ;M .: M. M. M. H. á G. G. G. G. G. G. R. R. X, n .: . ; Jadraque, Lora, de Tenthredinidae sp.; Empididae. Muscidae; phidae: udljr,Checa, Guadalajara, sp. s.n Garden, Kirstenbosch althaeoides: P. in cult. AFRICA, SOUTH 178, Narubis, Keetmanshop, MIBIA, n Alarc and sp., lecta Eulalia, (MA-591592); Pass, (MA). Anenous Garden Kirstenbosch in cult. AFRICA, SOUTH 4930, X, Karibib, Namibrand NAMIBIA, 257, X, IA aeB Esperanza, B. Cape RICA, teatus tera: Aldasoro nium (MA); gidum Garden, Kirstenbosch in cult. AFRICA, Alarc SOUTH and 652, Slankopf, X*, AFRICA, Jacq.: SOUTH n, X*, Kipplantivier, AFRICA, SOUTH Gardens, Kew num Cape, AFRICA, SOUTH (MA); Plaaf, Barus AFRICA, SOUTH 696, X, s.n. dae. Norfeu, .minimum Namaqualand, AFRICA, P. SOUTH 390, X, Sweet: (Andr.) Garden, Kirstenbosch Ermenek-Anamur, TURKEY, 1232, X*, Garden, Fenzl.: Stellenbosch at cult. RICA, Pomona, AFRICA, Garden, Kirstenbosch in Pass, cult. Franshoek AFRICA, SOUTH n, 9092 Aldasoro (COI). Cape, A Garden, Owan Mae oides Hymenoptera dae: phoridae: stoma (P); Inhaca, da carnatum Ilha MOZAMBIQUE, n, A-F, Yzerfontein-Darling, il. ,40 OT FIA aeB Esperanza, B. tetragonum Cape P. AFRICA, SOUTH 480, X, Garden, Willd.: Kirstenbosch in cult. 6, Moriscos, de Castellanos manca, Lepidoptera Bombylidae. and didae, visitors. unknown), .0,4, 0.408, of number ( total min data, per visits sample taxa, or visitor reference taxon, information: following eagnu acraeum Pelargonium A rdu alpinum Erodium (MA); . Diptera: .cicutarium E. M) Paarlberg, (MA); PPENDIX .inodorum P. Macropis p;Halictidae: sp.; Diptera: L)L (L.) Coelioxis alpoia;Msia;Ppnuia;Syrphidae: Pipunculidae; Muscidae; Calliphoridae; Knuth , – sp aailssn P;P whytei: P. (P); s.n. Cavanilles .reniforme P. L)L (L.) .glutinosum P. ypu ribesii Syrphus pi,Madrid, Spain, ypu ribesii Syrphus laoo9244 Aldasoro . laooadAlarc and Aldasoro , Coleoptera: laooadAlarc and Aldasoro 25 una Hontecillas, Cuenca, 9245; ó ó (L ypu ribesii Syrphus laooadAlarc and Aldasoro ’ 306 n Anthophora 9244 n dsr s.n. ldasoro ei. * 1,SUHARC,cl.i isebshGarden, Kirstenbosch in cult. AFRICA, SOUTH 718, X*, Herit.: .aridum P. (P); ’ : alpoia;Msia;Ppnuia;Syrphidae: Pipunculidae; Muscidae; Calliphoridae; .Pliaosadvstr fGrnaee epoiethe provide We Geraniaceae. of visitors and Pollinators 3. ei. onh:X 6,SUHARC,Cape, AFRICA, SOUTH 460, X, Moench.: Herit.) Cv)Wld:AF,n OT FIA WCape, SW AFRICA, SOUTH n, A-F*, Willd.: (Cav.) : ’ ei. * 1,SUHARC,Nuwerus, AFRICA, SOUTH 910, X*, Herit.: -* 6,SUHARC,Cp,Citrusdal, Cape, AFRICA, SOUTH 163, A-F*, ,n OT FIA uta tlebshGarden, Stellenbosch cult.at AFRICA, SOUTH n, X, sp.; Lf)L (L.f.) yande ayia,Zygaenidae. Satyridae, Lycaenidae, Bombylidae: p,Vespidae: sp., Andrenidae: : .australe P. eo3837 Aedo (MA); .scabrum P. laoo9040 Aldasoro il. ,28 AUSTRALIA, 228, A, Willd.: (MA); – Osmia M) an .Almad P. Jaen, (MA); Ad. Curt. (Andr.) – pi,Slmna atlao eMoriscos, de Castellanos Salamanca, Spain, ; itrs.n. Dinter Halictus ’ . Aldasoro Torymidae. ei. * ,SUHARC,cl.i Kirstenbosch in cult. AFRICA, SOUTH n, X*, Herit.: .A Dyer A. R. tl,L Italy, Jc. L (Jacq.) rg 26(P) 1276 Drege rcia,Malachidae. Bruchidae, (MA); , p,Crsde Halictidae: Chrysidae; sp., M) 6 .5,6, 0.153, 16, (MA), .caucalifolium P. laoo9025 Aldasoro Hymenoptera laoo9096 Aldasoro Sphaerophoria laoo9249 Aldasoro .elongatum P. . .A yr * ,SUHARC,Steinkopf, AFRICA, SOUTH n, X*, Dyer: A. R. Hymenoptera ,18 AUSTRALIA, 108, A, : sp., Hymenoptera rses.n. Krasse (MA); ó Hymenoptera aailss.n. Cavanilles Br.f)L f.) (Burm. ó 9233 n oblu maximus Bombylium — 81 9265 8831, n ó .triste P. p;Ihemnde Megachilidae: Ichneumonidae; sp.; ’ Dasypoda Andrena 8834 n Polistes qia Abruzzo, Aquila, odt) ieo bevto nhus( hours in observation of time data), no : ’ and (MA); ei. * 2,SUHARC,cl.at cult. AFRICA, SOUTH 526, X*, Herit.: laoos.n. Aldasoro (MA); .cortussifolium P. laoo9095 Aldasoro : : Aldasoro ab s.n. Tambs laooadAlarc and Aldasoro ,31 OT FIA Fiahamstown, AFRICA, SOUTH 361, X, .daucoides E. ,n ut nMdi Garden, Madrid in cult. n, X, M) ,00,4, 0.09, 5, (MA), ,80 MOZAMBIQUE, 860, X, Alarc M,P); (MA, ; M) rnd,M.Ca Mt. Granada, (MA); laoo9096 Aldasoro L)L (L.) laoos.n Aldasoro p Bombylidae: sp. (MA); nrnde;Anthophoridae: ; Andrenidae : (MA); Cv)Salisb. (Cav.) sp. .myrrhifolium P. M) aaoa oiad Arag de Molina Zaragoza, (MA); .alchemilloides P. laoo9241 Aldasoro lctr9139 Slechter p,Halictidae: sp., nrnde Halictidae: Andrenidae; : .crithmifolium P. te s.n. Strey sp.; ’ aq:X,43 OT AFRICA, SOUTH 483, X*, Jacq.: Andrenidae: : edl2947 Seydel ei. * 1,SUHAFRICA, SOUTH 315, X*, Herit.: ó M) udljr,P Guadalajara, (MA); (MA); .chium E. n nhpoia,Cymbicidae. Anthophoridae, : Coleoptera: é ..(MA); s.n. .Ri or 738 Moura Reis A. 22 ,01,8, 0.13, 9, 9282, , ’ n, Ceratina ei. ,n OT AFRICA, SOUTH n, X, Herit.: (P); .grossularioides P. .chamaedrifolium P. (MA); .trilobata M. Aldasoro – Diptera: Rechinger (MA); laoo9148 Aldasoro Syrphidae: .cucullatum P. .ovale P. eoe l 8256 al. et Aedo L Rechinger pi,Bro,Sargentes Burgos, Spain, CI;SUHAFRICA, SOUTH (COI); (MA); . ’ Halictus ei. *,65 SOUTH 645, , X* Herit.: Coleoptera: – : .botrys E. (COI); ó (COI); (MA); p;Vespidae: sp.; .cotyledonis: P. 9250 n * 2,SUHAF- SOUTH 223, X*, ,60 OT AF- SOUTH 610, X, : .umbellata M. pi,Grn,Cabo Gerona, Spain, M) eul Santa Teruel, (MA); aailss.n Cavanilles laoo9091 Aldasoro .stipulaceum P. oblu maximus Bombylius Andrena Oedemeridae. Halictus L)L (L.) and .E Sm. E. J. Br.. L (Burm.f.) ypia,Empi- Syrphidae, esMoura Reis (MA); .endlicherianum P. es * ,NA- n, X*, Kers: .pseudofumari- P. .vandierietiae M. (MA); .denticulatum P. p;Megachili- sp.; psrhsbal- Episyrphus M) 1 0.05, 11, (MA), ypu ribesii. Syrphus Alarc ñ – laoo9035 Aldasoro Mendon Diptera: ’ L)L (L.) ei. F-A*, Herit.: (MA); L)L (L.) adillas laoos.n Aldasoro (MA); pi,Sala- Spain, p,Antho- sp., á rg 1838 Drege sp. Oedemeri- .luridum P. mcsde lmaces : M) 10, (MA), Megachile aq X*, Jacq: ó .betuli- P. eosn. Aedo Aldasoro Aldasoro Aldasoro Aldasoro ,1100, X, Harvey: n Halictus Melano- ut in cult. (MA); . Polistes ’ ’ ’ .cico- E. 9262; , Herit. Herit.: Herit.: (MA); (MA); .ful- P. ç .in- P. Alda- s.n. a Dip- (L.f.) Syr- Me- — ó n : : . ; : Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ia,Cletde nhpoia,Adeia,Apidae: Andrenidae, and Anthophoridae, Collettidae, tidae, caespitosum G. mendica. drenidae: 0 .8,4, 0.383, 10, austroapenninum G. Fringillidae. noptera: caenidae. Apidae: Oedemeridae. deae, pi,Slmna atlao eMoriscos, de Castellanos Salamanca, Spain, U 4, 0.167, 4, (MA), cinereum 4, (1985): sp. leoptera: Anamur, and Ermenek between Satyridae: dae. Andrenidae: Pentatomidae. dae. dae: drena sp., Psithyus Muscidae. mellifera Apis iia;Payeia,Clihrde ucde Syrphidae sii Muscidae; Calliphoridae, Platypecidae, pididae; dae: Syrphidae: M) 5 .9,7, 0.194, 15, (MA), Ceratina 6, Espu de Sierra Murcia, (MA); 29(A;Gaaaaa Ca Guadalajara, (MA); 9239 Alarc Lepidoptera drenidae: dae: .8 4, 0.08, 9216 8825, litta sp. Scaeva Alarc and soro bollas, drenidae: Portillo, del Puerto Valdecebollas, Palencia, (2000), al. Sphecidae. Platigasteridae, drena scripta. Sphaerophoria Syrphidae: albomaculata, Scaeva Tachinidae, Phoridae, cidae, Norfeu, Cabo Lasioglosum 341 Oedemeridae: GERANIACEAE OF BIOGEOGRAPHY tera: & PHYLOGENY AL. ET FIZ 9274 9243 9263 9213, Eumenes : Alarc and soro 2008] Asilidae: s.n. Bernardos Moulares, Tunisia, Halictidae: eroptera Miarus Halictidae: mus balteatus aertadHcihn erCotE Fejaj, El 6, Chott 0.124, near 76, Hachichina and km Tamezret Douz, and Matmata Matameurtra, after Matmata, and dedine ñ . eaimarboreum Geranium Coleoptera: a, Hymenoptera . Bombus sp. Andrena Halictus Oedemeridae. ó psmellifera Apis Hymenoptera nhpoia,Friia,Halictidae: Formicidae, Anthophoridae, , M) an ioAlmad Pico Jaen, (MA); Diptera: laooadAlarc and Aldasoro M) uca irad Espu de Sierra Murcia, (MA), 3472 n sp. Aldasoro sp.. , Diptera: Miridae: : sp. .macrocalyx E. psmellifera Apis , Apidae: Coleoptera: – — Dasytidae. Sphaerophoria Asilus Andrena sp .caespitosum G. M) una Masegosa, Cuenca, (MA); Hymenoptera Andrena nrn;Hpii lirn,Macropis, albifrons, Hoplitis Andrena; Apidae: M)Tre,Pet eCabigordo, de Puerto Teruel, (MA) Diptera: Halictus pi,Snadr Castrovalnera, Santander, Spain, ypu iei,Eiipu balteatus. Episirphus ribesii, Syrphus ,8 Coleoptera: Lassioglossum .paularense E. Hymenoptera sp., .foetidum E. . : spp. Halictidae: . Sphecidae ó M) una Hontecillas, Cuenca, (MA); sp, sp Andrena Lycaenidae SL) Ghomrassen, (SALA); ó laooadAlarc and Aldasoro – 8828 n .maculatum G. ypia,Conopidae. Syrphidae, 9217 n Oedemera 32, Coenonympha ., Buprestidae. and Doufurea p,Sahpaia,Anthomiidae. Scathophagidae, sp., Diptera: Lassioglossum nhpoia,Halictidae: Anthophoridae, ypia : Syrphidae Anthophoridae: : .caespitosum G. psmellifera Apis Halictidae: , sp oepea Hemiptera. Coleoptera. nrnde:Adeas. ybcde Formicidae; Cymbicidae; sp.; Andrena : Andrenidae : laooadAlarc and Aldasoro Coleoptera: Aldasoro Diptera: sp psmellifera Apis Amirida sp. Syrphidae: Alarc .; Diptera: Diptera: M) ,01,12, 0.12, 8, (MA), ., Chrysomelidae: ybcde Formicidae. Cymbicidae, , – sp. M) 1 .1 6, 0.11, 11, (MA), – ,Lassioglosum Halictidae: ó , sp., – Chrysidae: 8627 n aahde otyhde Curculionidae Bostrychidae, Malachidae, ybcde Andrenidae: Cymbicidae, : – eersadS on(98:1, (1988): John St and Medeiros Sapygidae pi,Cec,Tragacete, Cuenca, Spain, – .malacoides E. . – p;Chrysomelidae: sp.; , ó esn (1989): Hessing p;Ichneumonidae: sp.; Muscidae. p Sphecidae: sp, Halictus pi,Tre,Javalambre, Teruel, Spain, n pi,Mdi,E Paular, El Madrid, Spain, Apidae: Anthophoridae: : orz(04:11, (2004): Goertz ypu ribesii Syrphus 03(A;btenTtoieadChenini, and Tataouine between (MA); 3043 sp. .glandulosum E. 82(A;Tre,Pet e Portillo, del Puerto Teruel, (MA); 8832 Bombylidae: ilo ta.(99:3, (1979): al. et Willson sp. ñ é ñ Diptera: ypia,Bombylidae: Syrphidae, n ucdeadEmpididae. and Muscidae Scaeva a, amares, , p,Sphecidae. sp., M) ,019 5, 0.189, 7, (MA): Halictus – enro s.n. Bernardos ypu iei,Sav p,Paragus sp., Scaeva ribesii, Syrphus ó Mordellidae obsfairn,B bifarius, B. flavifrons, Bombus Hymenoptera laooadAlarc and Aldasoro Neuroptera. 23 9224 9253, n laooadAlarc and Aldasoro – tl,Abruzzo, Italy, laooe l 9150 al. et Aldasoro Megachilidae: , n Halictidae: and laooadAlarc and Aldasoro Halictus laat (1980): Pleasants . laooadAlarc and Aldasoro psmliea Bombus mellifera, Apis Chrysis ñ sp Anthophora Lepidoptera Hymenoptera: enro s.n. Bernardos ó a, p,Muscidae. sp., 8828 n Diptera: . Lachnaia pde: Apidae ; Bombylidae: Hymenoptera: laooadAlarc and Aldasoro – sp. laooadAlarc and Aldasoro Hymenoptera Diptera: laooadAlarc and Aldasoro enro s.n. Bernardos Bombylius Stigmus pi,Salamanca, Spain, — Bombylidae: , sp., ypu ribesii Syrphus Formicidae. .collinum G. , , p,Halictidae: sp., – M) 10, (MA), M) 7 .7,36, 0.178, 17, (MA), — Dermestidae. .pelargoniflorum E. laooadAlarc and Aldasoro Halictus — laooadAlarc and Aldasoro Lasioglosum Formicidae: : Clytra .glandulosum E. pi,Plni,Valdece- Palencia, Spain, Anthophora SL) ewe Djebel between (SALA); laooadAlarc and Aldasoro Ryssa enro s.n. Bernardos Syrphidae: ,6, Halictus p,Anobiidae. sp., Hymenoptera , Megachilidae: sp., Coleoptera: err ta.2073 al. et Herrero : psmellifera; Apis — Megachile sp. ó ó Lycaenidae: Halictus .brevicaule G. Syrphidae: SL) ewe Me- between (SALA); Hymenoptera: 20 9259 9220, n — 9233 n ó sp nhpoia,Api- Anthophoridae, , ó laooadAlarc and Aldasoro Andrena — 82( 8832 n laooadAlarc and Aldasoro laooadAlarc and Aldasoro oblu maximus, Bombylius Eucera 84 9214 8824, n M) ,02 3, 0.2, 9, (MA), sp. Hymenoptera Hymenoptera Hymenoptera p;Curculionidae: sp.; , . p,Mgciia : Megachilidae sp., .glaucophyllum E. — miia,Mus- Empididae, , Bombylius , — Andrenidae: : SL).between (SALA);. Ichneumonidae, , Andrenidae: – Lepidoptera: , — , oblu maxi- Bombylius , Melittidae: — , pi,Cuenca, Spain, Hymenoptera: Hymenoptera p,Sphecidae. sp., Heteroptera: . M) ,0.142, 9, (MA), ,3 ypu ribesii, Syrphus ó ó sp psmellifera Apis — Volucella ó Diptera: 20 9231, 9210, n p,Apidae: sp., 85 9246 8835, n , A;Teruel, MA); p,Formici- sp., ypu ribe- Syrphus p,Apidae: sp., 25 9240 9255, n Halictus laooand Aldasoro Formica – Coleoptera: . Chrisomeli- SL) 12, (SALA), sp Diptera. , – 12, Andreni- : Episyrphus .molle G. — – – Lyssandra . Megachile Vespidae Gerona, ; io et Simon ó Halicti- , Diptera: Coleop- ó sp p,and sp., M) 8, (MA), ,9235, n, ó Turkey, , Birds: , Philipp Hyme- 9213, n 9201 n Meloi- Halic- (MA), . An- : An- : An- : Alda- Het- Em- Me- An- Co- An- sp., sp sp., Ly- ó ó ó G. n n n – – . : , , ´ salmoniflora Bitterfontein, prosopa l M) . mo Garoeb Formicidae of Km 1.5 (MA); eehde: Gelechidae Formicidae: Hylaeus Exoprosopa 0k oNuwerus, to km 20 nrnde eahlde Apidae: yesoense Megachilidae, G. Andrenidae, occidentales. 7, B. (2003): bifarius, Washitani B. rufocinctus, Bombus rau,Eiat uciols odlaatrata Mordella Dialictus puncticollis, Epicauta ornatus, irsrpe ieramh,Lmie otcs anr utt,Cephon guttata, Parnara viscossisimum boeticus, Lampides G. maha, hylas. Zizeeria rapae, Pieris 30, 0.209, 27, (2002): i,M eoa hlcdm psua eaiaiaa,Ceratina iwatai, chinensis Ceratina Polistes spissula, Chalicodoma remota, M. sis, chilidae: Halictidae: Formicidae; sp. h Paragus ferquens, rhous; Metasyrphus balteatus, Episyrphus macrogaster, ria cus. Anthomyiidae: Syrphidae: Hymenoptera: 35 0.059, 7, (1993): phidae ictidae: Hymenoptera: poo, Andrenidae. Halictidae, Hymenoptera: ypu ribesii Syrphus exiguous dius ard lEscorial, El Madrid, sp. Bombus mellifera; Apis Apidae: Muscidae. Coleoptera: melifera. Apis farius, Scathophagidae. didae; 9248 asn(00:1, (2000): Hansen sp. Halictus Alarc and robertianum G. acanthidactyla. A. punctidactyla, sinapis Leptidea damines, sp. Osmia phaeoptera Stelis tum, menoptera: dae: tris u assimilis nus richardsonii G. hypocrita. Syrphidae: : Halictidae Hymenoptera: ictidae: ribesii c Pieridae tera: cum Megachilidae: Collettidae; sp.; Apidae: pratense G. sp. bus 2, (2000): al. et Dlussky Andrenidae: menoptera: basalis Phaonia pipiens Syritta Syrphidae heritieri ó 8821 n osnaciliata Monsonia Empididae: ; Lepidoptera – Halictus M) ,023 ,Syrphidae 6, 0.233, 6, (MA), ypia : Syrphidae laooadAlarc and Aldasoro ; Calliphoridae: pi,Santander, Spain, hni campestris Rhingia : eaotm sp. Melanostoma p;Muscidae. sp.; aitsclatm .pratense G. calcaetum. Halictus irsnp.G phaeum G. napi. Pieris : Halictus Anthophoridae: ; p;Megachilidae: sp.; psmellifera Apis oloy eetaa .ynns .auiaa eahl tsurugen- Megachile acuminata, C. yanonis, C. fenestrata, Coelioxys ó M) ,003 4, 0.083, 5, (MA): – 9201 n parpoi sp. Sphaerophoria : Halictidae: Hemiptera esrcapensis Messor : Megachilidae: , sp. Chrysotoxum ltcersmanicatus Platycheirus – ot fia Springbock, Africa, South , Megachilidae: Halictus eiia;Btrde Curculionidae: Byturidae; Meliridae; Halictidae: Myrmicaria Staphilinidae: ; arpsfulvipes. Macropis Gelechia odlaatrata Mordella sp. , Empididae: ; laoo9078 Aldasoro Antomydae: , ot fia Aus-Wiputz, Africa, South nrnde Apidae: Andrenidae, Hymenoptera psrhsbalteatus Episyrphus Apidae: Halictidae: eolt aestiva Pegoplata Vespidae: Eumenidae: , Lycaenidae: : Empis p,Cleia,Megachilidae. Colletidae, sp., .palustre G. (MA) – – psrhsblets hni campestris. Rhyngia balteatus; Episyrphus 40, – — ot fia Steinkopf-Violdrof, Africa, South Lepidoptera. laoo9035 Aldasoro — , Diptera: p Colletidae. ; sp. laoos.n. Aldasoro tmriaobsoleta Stomorhina pi,Cl nMA, in Cult Spain, Scolidae: ; hlsoacmauau,C ircr,Ceiyssp.; Coelioyis nigricorn, C. campanularum, Chelostoma ,Apidae: 5, , sp. 70, , . Diptera: aitscalcaetum Halictus ypu ribesii Syrphus : Hymenoptera: — p;Msia.Andrenidae: Muscidae. sp.; Hymenoptera: – , Hymenoptera Hymenoptera: – ó obsdvru,Ai cerana Apis diversus, Bombus Dialictus Alarc sp., .subargenteum G. sp 9196 n irsnp,P rapae P. napi, Pieris M , laoo9068 Aldasoro , re 17) ,002 56, 0.012, 9, (1978): Green re 17) 9, (1978): Green Diptera: hlsoasp., Chelostoma .rostrata R. — oits Eumenes Polistes, nhpoaterminalis Anthophora Empis . Halictus aigosmsiuu,Lasioglossum scitulum, Lasioglossum Hoplitis . hlsoas.G sanguineum G. sp. Chelostoma M) ,003 4, 0.033, 3, (MA), Phorbia , . Diptera: Anthobium – Hymenoptera: , Crematogaster , drudeana Anthophoridae: : ó Syrphidae: rcoe ornatus Trichodes Lepidoptera Hymenoptera: Anthidium n laoo9261 Aldasoro and n eohlspnuu,Eyypu balteatus Epysyrphus pendulus, Helophilus Lepidoptera: lsk ta.(00:16, (2000): al. et Dlussky luiaorbitulus Albulina uee samuray Eumenes – Muscidae: . Scolia ugra tVitosha, Mt Bulgaria, Hymenoptera: M) ,021 6, 0.211, 7, (MA), parpoi sp. Sphaerophoria : .robertianum G. p;Agromyzidae: sp.; Lepidoptera: M) ,025 6, 0.275, 9, (MA), Dolichopodidae: ; (MA), sp Empididae: ; Apidae: , Syrphidae. Bombus p,Calliphoridae: sp., p;Vsia Masarinae: : Vespidae sp.; . obssp. Bombus , Apidae: ; psmliea obspratorum; Bombus mellifera, Apis .reinii G. psrhsbalteatus Epysyrphus Anthophoridae: : alpoia;Syrphidae: Calliphoridae; ; – p,Apidae: sp., Megachilidae: – Phasiidae: ; Megachilidae: Andrenidae: Sphaerophoria M) ,005 6, 0.005, 2, (MA), sp. aii,Garoeb, Namibia, Diptera: Andrenidae: , eaypu eais Sphaeropho- serarius, Betasyrphus sp., rco ta.(96:2, (1996): al. et Proctor – Apidae: laoo9887 Aldasoro laoo9036 Aldasoro laoo9049 Aldasoro hiosaculeipes Thricops : sp. — pi,Snadr loCam- Alto Santander, Spain, Diptera: eahlde Collettidae, Megachilidae, , sp. Gelechidae: Apidae: Hesperidae Hymenoptera: . Pterophoridae: ; – oloy nri.G pyrenai- G. inermis. Coelioxys psmliea .pratense G. mellifera. Apis Apidae: 56, , – Diptera: , pi,Santander, Spain, Hymenoptera: Lepidoptera obsrfcnts .bi- B. rufocinctus, Bombus Pieridae: Amegilla uoo(96:3, (1986): Yumoto pcuapuncticollis Epicauta Coleoptera. . .sanguineum G. . Apidae: ; – .sylvaticum G. Apidae: mi livida Empis .thunbergii G. ooia;Syrphidae. Conopidae; ioisborealis Limobius Lepidoptera ot 20) 35, (2004): Tofts obssp., Bombus M) ,013 5, 0.133, 5, (MA): , Coleoptera: Diptera: ynsm rotundata. Gymnosoma psmliea Bombus mellifera, Apis Coleoptera: Colletidae: . eczn sp. Leucozona Diptera: Andrena goyanigrescens Agromyza nrn sp. Andrena eioiia,Syr- Cecidomiiidae, laoo9038 Aldasoro oihpsplumipes Dolichopus sp., obsbaioa B. beaticola, Bombus Chelostoma psmliea Bom- mellifera, Apis nhdu manica- Anthidium eid biseta Melinda laooadAlar- and Aldasoro Bombylidae: nrn bicolour; Andrena eprscomma, Hesperis : M) ,00,4, 0.08, 6, (MA): M) 5K of Km 55 (MA); Amegilla p,Collettidae: sp., — Gelechia obslapponi- Bombus nhcai car- Anthocharis hni campes- Rhingia , M) Garies- (MA); Hymenoptera – – yit pipiens Syritta Megachilidae, p,Vespidae: sp., psmellifera, Apis , , laoo9065 Aldasoro : aaoand Nakano T Bombylidae: Halictidae: ; ;Muscidae: .; hlp and Philipp — Lycaenidae . Amblyptilia Halictidae, Halictidae, Halictidae: Syrphidae. Pieridae : – nigritellus – p Mega- sp. , Celonites Trichodes Halicti- ; Trichioti- – Diptera: Kandori Aldasoro p Hal- sp; Totland Syrphus Hylaeus Empi- ; ; sp. sp. — — Spain, — aemor- (MA); Zacla- Hy- . . Dip- Hal- , Exo- 70, , , odes Hy- — — M. M. – : ; , , , , , , ; : Delivered by Publishing Technology to: CSIC - REAL JARDIN BOTANICO IP: 161.111.170.152 on: Tue, 30 Apr 2013 08:02:49 Copyright (c) American Society for Plant Taxonomists. All rights reserved. om nSrc 19) 1, (1997): Struck in comm. aii hlba .fruticosum P. chalybea. tarinia 9148 Aldasoro elongatum P. peringueyi. Prosoeca Apidae: Nectariniidae Passeriformes, flabellifolium P. nus. Diptera echinatum P. lifera. dolomiticum nosum 2, (1997): Struck Nymphalidae: Lepidoptera: 1, (1995): blatt Diptera denticulatum P. rostrata. Philoliche bcrcp.P cucullatum P. obscuriceps. 19) 2, (1997): 4 tera crassicaule P. eyi. sp. odlt 19,19) 2, 1996): (1995, Goldblatt 2, (1997): Struck in Johnson carneum P. wahlembergiae. C. bergewhaliae, Prosoeca 5, (1997): Apidae: Hymenoptera 19) 3, (1997): Tabanidae: — Diptera Tabanidae: articulatum P. longirrostris. torhynchus 1, (1997): Goldblatt and Manning rostrata Philoliche a e at(90:1, (1990): Walt der Van Anthophoridae: menoptera 4 YTMTCBTN Vlm 33 [Volume BOTANY SYSTEMATIC Aus, 342 – 15, ,4 eagnu alchemilloides Pelargonium Nemestrinidae: : .coronopifolium P. – laoo9054 Aldasoro Hymenoptera 15, – : Tabanidae: : Tabanidae: : ot fia ut tKrtnoc Garden, Kirstenbosch at cult. Africa, South psmellifera Apis hllcersrt.P endlicherianum P. rostrata. Philoliche psmellifera Apis sp. Diptera — — — aitde Anthophoridae: Halictidae, : hllcersrt.P barkly P. rostrata. Philoliche ,4 hllcersrt.P betulinum P. rostrata. Philoliche – ,4 ,4 Hymenoptera itmn(93:1 .,8, 2.2, 1, (1993): Zietsman M) 1, (MA): Apidae: : – 15, – – — 15, 15, . mglaniveata Amegilla – Bombylidae: : — .alternans P. , Diptera ann n odlt 19,19) 1, 1996): (1995, Goldblatt and Manning – ,4 – M) fAus, of S (MA); Anthophoridae: : — Hymenoptera Hymenoptera hllceltrls .burgerianum P. lateralis. Philoliche hllcegls.P dipetalum P. gulosa. Philoliche rseaprnuy.P crispum P. peringueyi. Prosoeca ag es om nSrc 19) 1, (1997): Struck in comm. pers. Maggs tuk(97:1, (1997): Struck – – . , . — 5 yeotr:Apidae: Hymenoptera: 15, Diptera: Diptera lespr.cm.i tuk(97:1, (1997): Struck in comm. pers. Albers Coleoptera: — – psmellifera Apis — ,4, : alt 10) 1, (1908): Marloth , Tabanidae: : etrnafamosa Nectarinia , — Megachilidae: : — – ascrscassus; Tarsocerus — Lepidoptera: — , – oe 15) 1, (1954): Vogel , – , — , Diptera eaapscpni.P candi P. capensis. Megapalpus Nemestrinidae: : tuk(97:5, (1997): Struck Diptera – tuk(97:1, (1997): Struck — . Tabanidae: Birds: , ann n odlt 19) 1, (1995): Goldblatt and Manning — – Anthophoridae: : .antidysenthericum P. Anthophoridae: : eiotr:Nymphalidae: Lepidoptera: , , a e ate l 19) 1, (1990): al. et Walt der Van laoo9055 Aldasoro — Scarabeidae Anthophora – – . — tuk(97:1, (1997): Struck Bombylidae. : , tuk(97:1, (1997): Struck hllcelateralis Philoliche Nemestrinidae: : .cortussifolium P. Hymenoptera Diptera Amegilla – ,4 asrfre,Nectariniidae Passeriformes, ann n odlt (1995); Goldblatt and Manning Satyridae: . – – — – Hoplitis .flabellifolium P. 15, hllceformosa Philoliche Lycaenidae. uky Ermenek-Anamur, Turkey, — tuk(97:1, (1997): Struck ,4 , . Diptera Nemestrinidae: : – — — .capitatum P. . .crithmifolium P. — sp. 15, rsealniens P. longipennis. Prosoeca psmellifera Apis – M) ,007 4, 0.017, 2, (MA), ,4 ,4 – , Vespidae: ; ann n Gold- and Manning Diptera laoo9096 Aldasoro Diptera ehpaci statili- Neohipparchia mglanvaa A. niveata, Amegilla tuk(97:1, (1997): Struck mglaspilostoma Amegilla – – .appendiculatum P. Apidae: : 15, – 5 Hymenoptera: 15, — — – Nemestrinidae: : Nemestrinidae: ; rseaperingu- Prosoeca tuk(97:1, (1997): Struck – ,4 ,4 cetm and Scheltema .glutinosum P. Hymenoptera — ann and Manning – cans – – — Tabanidae: : Tabanidae: : lespers. Albers 15, ,4 15, , , – .gulosa P. . — psmel- Apis — – Celonites Diptera Diptera 20, – – Princeps .gluti- P. Moegis- Birds: , — ,4 Struck — Struck Struck — (MA): : Dip- , Nec- , – , Hy- 15, — — — — – – . , ; : , , , : : capensis. trifoliatum capensis 3, palpus (1992): Walt der Van and McDonald menoptera eigei .spinosum P. peringueyi. 1, (1995) aess Lithorina capensis, docus ogpni.P praemorsum P. longipennis. 4, dae. (1990): al. et Walt der Van Hymenoptera dae: 1, 1996) (1995, Goldblatt and ning Hymenoptera 4 — Diptera magenteum P. discoidea. ot fia ut tKrtnoc Garden, Kirstenbosch at cult. Africa, South Nemestrinidae scabrum P. spilostoma. 2, (1997): Hymenoptera 4 9027 al. et Aldasoro Bombylidae: mglaniveata. Amegilla hllcefroa .gls.P pilosellifolium P. gulosa. P. formosa, Philoliche luteolum P. phaleratus. Anastoechus 1, (1997): blatt mosa oti.P longiflorum P. rostris. laevigatum 1, — (1997): Struck in estrinidae: comm. pers. Steiner (1995), ni.P incrassatum P. ensis. — phyllum 1, (1997): Struck – – 5 itr:Bombylidae. Diptera: 15, , , 5 oepea Scaraboidea: Coleoptera: 15, , — — — , .papilionaceum P. . aiu uhnts ytehssarroti.P rapaceum P. scabrirrostris. Systoechus eurhynatus, Parisus yeotr:Vsia:Masarinidae: Vespidae: Hymenoptera: , , .gls.P longicaule P. gulosa. P. ,Hmnpea eiotr:Lycaenidae. Lepidoptera: Hymenoptera: 4, , .sericifolium P. Hymenoptera Nemestrinidae: : – — — tuk(97:1, (1997): Struck – – Anthophoridae: : rseaprnuy.P klinghardtense P. peringueyi. Prosoeca ,4 ,4 tuk(97:2, (1997): Struck tuk(97,1, (1997), Struck ihrn arpeu.P luteolum P. macropterus. Lithorina Anthophoridae : Apidae: : Anthophoridae: : Calliphoridae: ; – — Hymenoptera . – 15, Lepidoptera 20, — ,4 p;Tabanidae: sp.; ,4 Hymenoptera – M) 1, (MA): 20, Diptera – Formicidae. : – – – – – 5 yeotr:Apidae: Hymenoptera: 15, odlt ta.(95,MnigadGoldblatt and Manning (1995), al. et Goldblatt – – ann n odlt 19) odlt tal. et Goldblatt (1996), Goldblatt and Manning tuk(97:1, (1997): Struck Diptera tuk(97:5, (1997): Struck psmellifera Apis lespr.cm.i tuk(97:1, (1997): Struck in comm. pers. Albers rseaprnuy.P minimum P. peringueyi. Prosoeca ann n odlt 19,19) 1, 1996): (1995, Goldblatt and Manning tukadVndrWl 19) 2, (1996): Walt der Van and Struck – — – tuk(97:2, (1997): Struck Hymenoptera eetiia:Nemestrinidae: Nemestrinidae: : : mglanivetata Amegilla , — odlt ta.(95;MnigadGold- and Manning (1995); al. et Goldblatt — Lycaenidae: omn ucpni,Rycmamclt.P. maculata. Rhyncomya fuscipennis, Cosmina . Apidae: : — — Nemestrinidae: : ,4 — ,4 Lepidoptera mglaspilostoma Amegilla eirci capicola. Peritrichia , , Anthophoridae: : — hllceangulata. Philoliche ,3, – Diptera — – .myrrhifolium P. – 15, , 15, — , tuk(97:2, (1997): Struck . — Diptera Diptera .peltatum P. Diptera , Diptera Diptera , — — psmellifera Apis — Bombylidae: : ayesmarshalli Cacyreus Megachilidae. : ,4 ,4 , laoo9095 Aldasoro : – Diptera — Jugurtia , – Papilionidae: – Bombylidae: : Bombylidae: : tuk(97:1, (1997): Struck Tabanidae: : – 15, 15, A ,4 Bombylidae: : – Nemestrinidae: : — esadGs 20) 2, (2006): Gess and Gess ogsohnhslongir- Moegistorhynchus . psmellifera Apis – ot fia Darling, Africa, South ,4 iesps .tricolor P. diversipes. – Diptera 15, Diptera – . mglaatrocincta Amegilla Diptera tuk(97:3, (1997): Struck .pinnatum P. Bombylidae: : Hymenoptera tuk(97:2, (1997): Struck – .grossularioides P. sp., 20, Diptera — Anthophoridae: , – ,4 eaapscap- Megapalpus oe 15) 1, (1954): Vogel Diptera Masarina M) 2, (MA): Bombylidae: : Bombylidae, : .ocellatum P. – hllcefor- Philoliche . aii demo- Papilio Tabanidae: : 15, .scabrum P. — Megapalpus Megapalpus Bombyli- : . Bombomya .hetero- P. , – — — Diptera Prosoeca Prosoeca — – Nem- : — ,4 Struck ,4 Mega- Api- : sp. Man- — , , – – Hy- , ,3, 15, 20, — — — A P. – – – – , , , . :