Cucumis Zambianus (Cucurbitaceae), a New Species from Northwestern Zambia Mark P

NCRPIS Publications and Papers North Central Regional Plant Introduction Station

10-2008 Cucumis zambianus (Cucurbitaceae), a New Species from Northwestern Zambia Mark P. Widrlechner United States Department of Agriculture, [email protected]

Joseph H. Kirkbride Jr. United States Department of Agriculture

Amanuel G. Ghebretinsae St. Louis University

Kathleen R. Reitsma United States Department of Agriculture

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Abstract During germplasm explorations within Zambia in 1984, seven Cucumis accessions were collected that could not be identified to species. Two of the accessions were studied in-depth. Based on phenotypic characters, they were closest to Cucumis pustulatus. In ITS analyses of all available Cucumis species and the accessions, the two accessions grouped with 100% bootstrap support in a clade comprising C. anguria, C. dipsaceus, C. insignis, and C. pustulatus. The ca cessions differed from these four Cucumis species by the following characters: plants pilose, male inflorescences paniculate with 6-30 flowers, staminate-flower calyx lobes linear, pistillate- flower pedicels long and cylindrical, and a genetic difference of 4 base pairs in their ITS sequences. Controlled crosses were made within and between the two accessions and between each of them and C. pustulatus. Fruits were produced from crosses made within and between the two accessions, but attempts involving C. pustulatus were unsuccessful. The two accessions are described as a new species, Cucumis zambianus. Six of the seven unidentified Cucumis collections from Zambia are identified as C. zambianus herein. They ew re all collected in the northwestern corner of Zambia, but the new species is also expected to occur in eastern Angola and southern Zaire.

Keywords Cucumis, Cucurbitaceae, ITS, new species, phylogeny, Zambia

Disciplines Agriculture | Horticulture | Plant Sciences

Comments This article is from Systematic Botany 33, no. 4 (2008): 732–738.

Rights Works produced by employees of the U.S. Government as part of their official duties are not copyrighted within the U.S. The onc tent of this document is not copyrighted.

This article is available at Iowa State University Digital Repository: http://lib.dr.iastate.edu/ncrpis_pubs/21 Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 19)monograph. (1993) Hlia n efe 98 rtems eetmngahof monograph recent the most genus the using the or 1978) Re- them Jeffrey and identify and 505597 (Halliday Widrlechner not PI interesting. accessions, could very these itsma were of 505608, Two PI Iowa. and Ames, Introduction in Plant Regional Station Central North USDA-ARS the genus to the Introduction crops. of individual accessions Plant of Sixteen conservation the and the to for number, responsible distributed Stations Introduction then Plant were a accessions Each assigned System. Germplasm was Plant accession National do- USDA were the exploration to that nated from The samples as 1984). seed (Attere Zambia, year, exploration in following plant samples IPGRI-funded germplasm an of of part number col- Zambia, large Chilanga, a of Station, Mlongoti lected Research G. Makulu and Mount Italy, the Rome, (IPGRI), Resources Genetic n I527 eeslce o hsproebcueteunknown the 343701 because PI purpose (interspecific). this 532673) for PI selected cumis and were 343701 532673 PI (PI and f. Hook. and ex 505608 or Naudin 505597 PI accessions between different and same of the (between-population), plants of two plants between two (within-population), between (self-pollination), accession plant same the from ers vines the at least and at maintained ensure photoperiod. planting were to 12-hr lighting after temperatures supplemental a night with d respectively, and 26°C, 10 Day and 29°C benches grew. they greenhouse as on trellised set plas- a 18.9-L in pots to transplanted planted tic were then Seedlings Zambian and mix. original soil germination greenhouse of facilitate standard samples to chipped stored were representing collections, 505608, PI and 505597 oyih 08b h mrcnSceyo ln Taxonomists Plant of Society American the by 2008 Copyright © Botany Systematic a o td,adi a ocue htteeaccessions genus these the that of concluded described. species here was new it Ghebretin- a to and represented material study, leaf for fresh and sae Kirkbride to sent were 2 olntoswr fetdb aduigsaiaeadpsilt flow- pistillate and staminate using hand by effected were Pollinations Crosses— Controlled n18,A .Atr fteItrainlBadfrPlant for Board International the of Attere F. A. 1984, In SA giutrlRsac evc,US ainlAbrtm lrl&NreyPat eerhUi,30 e York New 3501 Unit, Research Plants Nursery & Floral Arboretum, National U.S. Service, Research Agricultural USDA, akP Widrlechner, P. Mark eei ifrneo aepisi hi T eune.Cnrle rse eemd ihnadbtentetoacsin n between and accessions as two identified the between are and Zambia a within from and made cylindrical, collections were and crosses Controlled long sequences. pedicels pustulatus ITS pistillate-flower and their linear, them in lobes of pairs calyx each base 4 staminate-flower of flowers, difference 6–30 genetic with paniculate inflorescences male dipsaceus C. pce sas xetdt cu nesenAgl n otenZaire. southern and Angola eastern in occur to expected also is species pce.Too h cesoswr tde ndph ae npeoyi hrces hywr lss to closest were they characters, phenotypic on Based available in-depth. all studied were of accessions the of Two species. cesoskydotms lsl to closely most out keyed accessions 1 uubtca,IS e pce,pyoey Zambia. phylogeny, species, new ITS, Cucurbitaceae, Keywords—Cucumis, uiggrpamepoain ihnZmi n18,seven 1984, in Zambia within explorations germplasm Abstract—During SA giutrlRsac evc,NrhCnrlRgoa ln nrdcinSain oaSaeUniversity, State Iowa Station, Introduction Plant Regional Central North Service, Research Agricultural USDA, uui zambianus Cucumis 3 an oi nvriy eateto ilg,30 ald vne t oi,Msor 30-00U.S.A. 63103-2010 Missouri Louis, St. Avenue, Laclede 3507 Biology, of Department University, Louis Saint 5 ot eta einlPatItouto tto,Iw tt nvriy ms oa50117 U.S.A. 50011-1170 Iowa Ames, University, State Iowa Station, Introduction Plant Regional Central North Cucumis 20) 34:p.732–738 pp. 33(4): (2008), eeuscesu.Tetoacsin r ecie sanwspecies, new a as described are accessions two The unsuccessful. were , .insignis, C. Cucumis M n1Mrh20,se ot f2 ed aho PI of each seeds 25 of coats seed 2006, March 1 On riswr rdcdfo rse aewti n ewe h w cesos u tepsinvolving attempts but accessions, two the between and within made crosses from produced were Fruits pustulatus. C. Krbie19) ebru specimens Herbarium 1993). (Kirkbride TRASAND ATERIALS pce n h cesos h w cesosgopdwt 0%bosrpspoti ld comprising clade a in support bootstrap 100% with grouped accessions two the accessions, the and species and 4 isuiBtnclGre,PO o 9,S.Lus isui616U.S.A. 63166 Missouri Louis, St. 299, Box P.O. Garden, Botanical Missouri 1,6 .pustulatus C. oehH ikrd,Jr., Kirkbride, H. Joseph 6 Cucumis uhrfrcrepnec ([email protected]) correspondence for Author M .pustulatus C. ETHODS Ccriaee,aNwSeisfo otwsenZambia Northwestern from Species New a (Cucurbitaceae), h cesosdfee rmteefour these from differed accessions The . vneN,Wsigo,D 00-98U.S.A. 20002-1958 DC Washington, NE, Avenue .zambianus C. .wr transferred were L. Cucumis omnctn dtr akP Simmons P. Mark Editor: Communicating lr Zambesiaca Flora sn Kirkbride’s using ms oa50117 U.S.A. 50011-1170 Iowa Ames, .pustulatus C. hc is which , een hywr l olce ntenrhetr onro aba u h new the but Zambia, of corner northwestern the in collected all were They herein. Cu- 2 mne .Ghebretinsae, G. Amanuel 732 .Sgn) h T einwsslce,a thspoe sfli clari- in Ghe- useful 1998; proven al. et has (Jobst it transcribed cucurbits as among internal selected, relationships was the phylogenetic region intervening fying amplify the ITS (including The to DNA gene). ribosomal used 5.8S nuclear of then region was (ITS) spacer DNA (Qia- specifi- Kit manufacturer’s The Mini following Plant cations. with California) sequencing. DNeasy bags Valencia, the DNA Corporation, plastic using for isolated gen sealable Ghebretinsae was in to DNA field-grown placed genomic overnight Total 505608, caged, shipped PI and from and gel, collected silica 505597 were PI of accession) plants per leaves 505608. young PI range and the 505597 within regeneration PI fell by any for if displayed greenhouse determine variation the to of and in verification samples taxonomic seed and original from as started only 2006. identified October exploration 6 Zambian on Her- collected 2006. were October accession 11 each on occurred of not queenright frost vouchers did 505608 killing small barium PI a a pollination. before for so cage flowers the 505597, develop in PI placed was on colony present bee honey were staminate both flowers mois- 2006, pistillate September preserve 11 and to On competition. newspaper weed al. chopped reduce and et with ture (Clark mulched diseases were insect-vectored with Plants from covered 1991). plants immediately the were protect frames to Cage screening temperature Zambia. and local in photoperiod when found September the regimen and parallel Ames, August closely in late most in would cages develop conditions field to them pollination 2006, allow into August to 10 Iowa, transplanted On were greenhouse. seed the original seedlings the in from pots these (also peat 505608 in PI started and were 505597 July samples) PI 26 of on each Zambia, seeds in 15 found un- 2006, those expressed resembling stably closely more were conditions conditions der greenhouse under were observed pepos acters open-pollinated discarded. All and number. vines the accession a from tagged and with removed was again date pollination secured Each the and contamination. with pistil external prevent the pistil. to around the clip closed onto hair brushed then was were pollen petals the and The anthers; staminate the corresponding expose five to flowers from flowers pistillate removed its were Undamaged exposed petals contamination. unclipped; have pollen were may to that following pistil damage or the for previous stamens checked the began was closed bud buds Pollinations Each staminate buds bud. afternoon. accession’s staminate an pistillate five collecting least by each morning at for accession, pistil- such an closed all within were After located clips. were metal buds small late with closed were morning lowing oeua Analysis— Molecular 1984 the from received accessions additional five 2006, October 26 On Study— Morphological Controlled fol- the open to were that buds floral pistillate afternoon, the During Cucumis i ftesvnunidentified seven the of Six zambianus. Cucumis Cucumis cesoswr olce htcudntb dniidto identified be not could that collected were accessions pce ytefloigcaatr:pat pilose, plants characters: following the by species n2 uut20,fehla ape (three samples leaf fresh 2006, August 21 On 3,4 uui pustulatus Cucumis n ahenR Reitsma R. Kathleen and ovrf htmrhlgclchar- morphological that verify To Cucumis p Apni )were 1) (Appendix sp. nISanalyses ITS In . .anguria, C. Cucumis C. 5 Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. evdi ie fP 068goni h il n green- PI of and morphology field floral the compare in to grown ob- unable 505608 were were PI We morphology house. of foliar vines field in in greenhouse. fall served the differences the in in significant grown grown those No 505597 to PI compared when of planting vines in observed were failed. crosses these ipeadto ftx,btsvn nyoete e replicate. per tree one and only saving swapping but branch taxa, TBR of using calcu- addition by was simple pseudoreplicates support 10,000 1985) from Felsenstein tree-bisection- lated (BS; Bootstrap the addition replicates. random used sequence 1,000 with searches option, branch-swapping char- Heuristic (TBR) all reconnection weighted. with and equally characters acters (MP) uninformative (Swofford parsimony parsimony 4.0b10 excluding maximum PAUP* in 2002), under conducted were performed analyses Parsimony were analysis. data of analyses combined Phylogenetic S1797). trees the by number resulting (study and obtained TreeBASE matrix in be data archived and can The are EU563019 (2007). sequences al. are other et 505608 Ghebretinsae Gen- of PI referencing that (2000). and and Ochoterena 505597 respectively, PI and EU563020, for Simmons the numbers following of accession characters, method Bank as coding coded were indel 2.0a11; appro- indels the simple (version and with codes, Se-Al coded IUPAC were in priate positions adjustments Ambiguous 1996–2002). manual Rambaut by followed Michigan) aetrsle n1 eo.W tepe 4cosswith crosses 24 attempted We female pepos. the as two 11 505608 in PI with Between- resulted parent crosses parent female respectively. and pepos, the pepos, 9 as in 505597 5 resulted PI and with pollinations 9 in population resulted of 505608 Self- production respectively. PI the pepos, of re- 8 pollinations and 505597 within-population 6 PI and of production of 505608. the PI in pollinations and sulted within-population 505597 the PI and both of for Self- Crosses successful 2006. all were July classes between-population 12 ma- and All on within-population, May. harvested self-pollination, into were continued and fruits planting) ture after d 12 (42 began April interspecific, and between-population, population, to prior 2006 October plantings 11 Field on development. 2006. frost fruit October killing a 6 by and destroyed 2006 were August 25 planting. on after ings longitu- d sliced 91 at fruit section mature cross a in with or vine dinally and of vine planting, portions to stem and after main tip, d crown, and 41 roots of staminate at true samples with representative beginning first vines flowers the and pistillate of present and portions still expanded, d 30 fully cotyledons at leaves plant with entire planting an including from growth, accession of each stages of various at taken were vouchers Multiple greenhouse. 0foee;crlalbso aefoesbodyaueat acute broadly flowers male of lobes corolla 3–10-flowered; racemose, 34’ inflorescences male rootstock; woody a without annual, Plants 34(14). in edited and assembled were data Con- combined recognition. the Sequencher™ species-level were to of deserve 2007), study sequences accessions al. this sensus two et from the (Ghebretinsae whether generated data test published data previously Sequence with Ghe- (2007). following combined done al. were et products PCR pu- bretinsae the amplification, of (PCR) sequencing reaction and chain rification Polymerase 2007). al. et bretinsae 2008] osgiiatdfeecsi oiro lrlmorphology floral or foliar in differences significant No adcosso l orcass efpliain within- self-pollination, classes: four all of crosses Hand plant- field the from collected were vouchers Herbarium the in rapidly grew 505608 PI and 505597 PI of Plants lnsana rprnil iho ihu od ottc;ml nlrsecspnclt,fsiuaeo oiayfoe;crlalbsof lobes corolla flower; solitary a or fasciculate paniculate, inflorescences male rootstock; woody a without or with perennial, or annual Plants .pustulatus C. litci uln,ots n urnt tae;ses4.8 seeds 8 apex; flowers at female mucronate of and pedicel obtuse apex; outline, at in mucronate elliptic and obtuse flowers male 5 seeds 1.5 apex; flowers at female of pedicel apex; vrin42 eeCdsCroainIc,AnArbor, Ann Inc., Corporation Codes Gene 4.2; (version cesos(I330 n I527) l of All 532673). PI and 343701 (PI accessions mln,2 long, mm –6 R ESULTS . mwd,adc.1m hc 6...... thick mm 1 ca. and wide, mm –2.5 mln;crlalbso eaefoesooaeo ral bvt notie ct rbodyacute broadly or acute outline, in obovate broadly or obovate flowers female of lobes corolla long; mm –7 IREHE TA. UUI ZAMBIANUS CUCUMIS AL.: ET WIDRLECHNER C ULT TO OUPLETS . mln,2.4 long, mm –6.7 S UPPLEMENT ln emls ytm cesbeoln thttp:// at online National accessible the www.ars-grin.gov/npgs. System, through Germplasm purposes Plant educational their and in of clarity research for Seeds originals here). the presentation from differences slight by with key species-level (2007; extensive Schaefer more current, a supplements pce-ee e to key Kirkbride supplements species-level set first The couplets. revised ( Arabia Saudi as east far ad ( lands 068wti the within 505608 species, described melo represent newly thus, the and, zambianus of 505608, PI populations and additional presented 505597 variation PI of accessions range the by and within 505607, fell PI clearly 505606, 505609) PI PI 505605, (PI Zambia from accessions fdsrbto ntoia atArc,rahn sfrsuhas south far ( as Africa center reaching Africa, a South east have tropical clade cross-compatible in this distribution the of of as Members 1993). to (Kirkbride referred group been within that traditionally resolved taxa have is African turn non-domesticated in comprising which clade clade, larger a this of consider member we the a 1), bp be Given to (Fig. 7 together 57%). of clade difference = this genetic uniting (BS a and low resemblance is morphological relationship this supporting dipsaceus containing clade a in 505608 4 morphological our a corroborating analyses. and 1), most (Fig. 100%), their taxa from = related grouping closely (BS acces- this support two separates strong difference the base-pair a that with showed grouped data sions the from Results of accessions. 505608 analysis PI parsimony and 505597 PI of status the 068bcuetefedpat aldt eeo lwr be- flowers frost. develop killing to the failed plants fore field the because 505608 yigteITS1 the lyzing i fteISdt atto received partition is data trees ITS port. these analy- the parsimony of of the (RI; consensus in sis nodes index strict Twenty-seven 1. retention The Fig. and in ensemble 0.748. shown an Kluge of and (CI; 1989) 0.516 index Farris 516 of consistency of 1969) ensemble trees Farris an parsimonious Parsimony with equally characters. steps 16,100 informative yielded additional (24.2%) analysis 63 197 re- which in characters as of sulted indels long, Scoring informative. characters parsimony 813 were was 747 matrix from data ranged region ITS 2 mln;crlalbso eaefoeselpi rbroadly or elliptic flowers female of lobes corolla long; mm –120 K hsnwseisi ecie eo,floigtost of sets two following below, described is species new This h ytmtcpsto facsin I559 n PI and 505597 PI accessions of position systematic The h T hlgn loapast lc I559 n PI and 505597 PI place to appears also phylogeny ITS The u td fISsqecsin sequences ITS of study Our ae npeoyi hrces oro the of four characters, phenotypic on Based IRKBRIDE . mwd,ad1.1 and wide, mm –3.2 L. .anguria C. and , seApni ) I551 a dniidas identified was 505610 PI 1). Appendix (see (1993) .anguria C. .anguria C. .SIS eesqecs h egho the of length The sequences. gene –5.8S-ITS2 dtrsnote: editor’s ,a a ot sCa ( Chad as north far as ), Cucumis Cucumis . mtik...... 35 ...... thick mm –1.8 Fg ) lhuhtebosrpvalue bootstrap the although 1), (Fig. ,a a eta h aeVreIs- Verde Cape the as west far as ), D .pustulatus C. ISCUSSION hlgn a siae yana- by estimated was phylogeny uui insignis Cucumis ohsplmnsaeformatted are supplements both 8 p h eutn aligned resulting The bp. –784 .zambianus C. es tit,adtesecond the and stricto, sensu Cucumis ). 0 otta sup- bootstrap Ն50% .pustulatus C. a mlctosfor implications has r vial for available are , .pustulatus C. .zambianus C. Cucumis ,adas and ), .anguria C. (1993) ’s 733 , sp. C. C. C. Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 734 Cucumis F IG 35’ 53) lnslcigwoyrosok efbae nrresrgs opls;ml nlrsecspnclt;pdclo eaefoes65 flowers female of pedicel paniculate; inflorescences male pilose; to antrorse-strigose blades leaf rootstock; woody lacking Plants 35(34). .Oeo h 2 otprioiu re bandfo h nlsso h T eune eutn rmprioyaaye fISdt of data ITS of analyses parsimony from resulting sequences ITS the of analysis the from obtained trees parsimonious most 324 the of One 1. . 3m og yatimo eaefoesantrorse-strigose flowers female of hypanthium long; mm 8–13 flowers female of pedicel scabrous; or antrorse-strigose blades Leaf 35a(35). 35a’ p. I559,P 068 n ugop,coe oilsrt rnhlnts otta values Bootstrap lengths. branch illustrate to chosen outgroups, and 505608, PI 505597, PI spp., lnsprnil ihawoyrosok efbae nrresrgs,saru,o ipdlu;ml nlrsecsfsiuaeo a or fasciculate inflorescences male hispidulous; or scabrous, antrorse-strigose, blades leaf rootstock; woody a with perennial, Plants oiayfoe;pdclo eaefoes8 flowers female of pedicel flower; solitary mln;futpdclpls ...... pilose pedicel fruit long; mm efbae ipdlu;pdclo eaefoes15 flowers female of pedicel hispidulous; blades Leaf ool oe ffml lwr 5.2 flowers female of lobes corolla 4 flowers female of lobes corolla outside; mwd;futpdclhsiuos...... 19...... hispidulous pedicel fruit wide; mm –8 5m og ri eie nrre rrtos-tioeo ipdlu 35a ...... hispidulous or retrorse-strigose or antrorse- pedicel fruit long; mm –45 . mwd;futpdclatos-o ers-tioe...... 20...... retrose-strigose or antrorse- pedicel fruit wide; mm –4.8 YTMTCBOTANY SYSTEMATIC 5m og yatimo eaefoespbrln rhsiuosoutside; hispidulous or puberulent flowers female of hypanthium long; mm –45 0 r hw bv h branches. the above shown are Ն50% .meeusei C. Vlm 33 [Volume .zambianus C. .pustulatus C. .Jeffrey C. –120 Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. .)m og oe o geniculate; not nodes long; aculeate; mm not (0.3–)1.52( –2.5) hairs, hairs nonbreakaway stem with pilose 5-sided, cate, nends(5.5 internodes monoecious. tubers, lacking ab- similis. cylindraceisque longis pedicellis femineorum florum bus, 6 floribus masculinis paniculatis, masculinis inflorescentiis pilosis, plantis eybodyoaet iclri uln,4 outline, in circular to ovate broadly very hairs, nonbreakaway with aculeate; pilose not petiole, each 6 on section, type single cross in sulcate drical, 57 ...... zambianus . . Cucumis ...... absent rootstock woody Annual, 36a. 2008] 6.Prnilwt od ottc 37 ...... rootstock woody with Perennial 36b. m (0.55 cm, rooaei uln,(2.5 elliptic outline, (rarely) in to obovate elliptic or broadly or entire pinnatifid, symmetrical, weakly to (rarely) asymmetrical 3 lobes outline, lateral apex; in the elliptic to 1.5 long, elliptic cm broadly to or ovate ovate broadly pinnatifid, narrowly twice or once symmetrical, sometimes pilose, lobe or central entire to hairs; antrorse-strigose nonbreakaway acu- below with not ciliolate intervenium pilose, the to the on antrorse-strigose on leate, and pilose, hispid to below antrorse-strigose veins surface upper truncate, the rarely obtuse, or cordate 0.5 sinus base basal the with serrate, weakly or entire eto,(6 section, sessile. buds, flowers flower branch a secondary with suppressed sometimes flower, 6 containing solitary paniculate, a female 3 male the fasciculate, sessile; the ones sessile; bisexual the flowered, aculeate. gynecandrous; not or simple, pilose, sexual solitary, long, tendrils apex; cm the 6–16(18.5) at obtuse or acute broadly u rcels yatimifniua ocampanulate, 2.8 to long, mm infundibular 3.5–5 hypanthium bracteoles; out ueuetisd;crlatb 0.6 tube corolla outside, inside; pilose puberulent sparsely infundibular, corolla pilose; at acute sparsely narrowly 2–3 base, wide apex, a to the flaring outline in linear 5–7, rbodyooaei uln,ots n urnt tthe at mucronate and obtuse 5 outline, apex, in obovate broadly or 5 lobes corolla diameter; in hu 1 thium Plants A 7.Ml nlrsec aiuae eie ffml lwr 65 flowers female of pedicel paniculate, inflorescence Male 57b. 7.Ml nlrsec aeoe eie ffml lwr 1.5 flowers female of pedicel racemose, inflorescence Male 57a. lnoia aua 3k fMiiug oKa- to G. Mwinilunga and of Attere S F. km A. 11 63 by mapanda, Kabuka, 505597) (PI at 1984 col- Mlongoti originally Jun seeds 16 from lected Iowa, Intro- Ames, nov. Plant Regional Station, sp. Central duction North Reitsma, USDA the R. at Grown K. & sae US). 2006, 572 May Reitsma 31 Zambia, Province, uuiinsigni Cucumi –10 eielt;pdcltrt rwal uct ncross in sulcate weakly or terete pedicel pedicellate; – eb,poal nul ihafbosrootstock, fibrous a with annual, probably herbs, ie ogrta ie ihtemargin the with wide, than longer times –)0.81 . maoeisbs;flmnsrdal com- radially filaments base; its above mm 1.6 4m og ioet nrresrgs,with- antrorse-strigose, to pilose long, mm –)814 × mwd,auet ral ct rots at obtuse or acute broadly to acute wide, cm –6 0 lrmmsuiou ayi oi lineari- lobis calycis masculinorum florum –30, blades 1m;saessprtn rmtehypan- the from separating stamens mm; 4–11 9 mlong. cm –)916(19) × Hltp:N;Ioye:B,IC ,M,P, MO, K, ISC, BR, Isotypes: NA; (Holotype: °54Ј mfaigt mwd tbase, at wide mm 2 to flaring mm 0.3–1 mde,adteae ral ct or acute broadly apex the and deep, cm –3 T . mi imtr ioe ay lobes calyx pilose; diameter, in mm –3.5 -o -amtl oe,bodyoaeto ovate broadly lobed, 5-palmately or 3- , AXONOMIC .pustulato C. ,24°8Ј S, irehe,J .Krb. Ghebretin- Kirkbr., H. J. Widrlechner, –)4.57.5 ,bodyelpi,bodyovate, broadly elliptic, broadly –7, Stems ,ee.130m Northwestern m, 1,320 elev. E, T REATMENT 3c og uecnea pubescence long, cm –13 , )m og 2 long, mm –1.5(3) .dipsaceo,etC. × rcmet lgtysul- slightly procumbent, Leaves IREHE TA. UUI ZAMBIANUS CUCUMIS AL.: ET WIDRLECHNER . m ct to acute cm, (1–)3.57.5 — irehe,Cak & Clark, Widrlechner, YE ...Iowa: U.S.A. TYPE: –11.5 – with C Inflorescences 10( ULT TO OUPLETS – 30)-flowered, × petioles 8–14(18.5) .anguria C. –5mm cylin- S Male uni- UPPLEMENT mln 14...... long mm –7 –5- 2 mln ...... long mm –120 –9 ln nrdcinSain ms oa rmsesoiial olce 16 collected originally seeds from Iowa, Ames, Station, Introduction Plant rse ncosscin 0.7 section, cross in pressed edd bogt bvt,irglrytioae bueto 1 apex, obtuse the trilobate, ex- at irregularly papillate connective and obovate, anther crenulate to edges; oblong the tended, along puberulent 1.5 long, sigmoid, free, thecae anther puberulent; minutely lbos icclnrcl 0.5 cylindrical, disc glabrous; hu ue oteoay ue oto ral ellipsoid, broadly portion fused hypan- 5 ovary; long, of mm the 3/4s 6–12 to to 2/3s fused lower thium ridged; hour-glass distinctly not hypanthium shaped, cylindrical; hairs, nonbreakaway 65– outline, glabrous. ameter, iebs,nrol ct tteae,2 apex, a the into at flaring acute outline narrowly in base, elliptic narrowly wide to linear calyx 5–7, inside; lobes puberulent aculeate, not outside, pilose sparsely oto apnlt,3.5 of campanulate, apex portion at 1/4 bristle upper base; hyaline tuberculate, 0.8 not aculeus long, mm 1.5–6 oe rnvreybodyoaeo litct ral elliptic 6 broadly outline, to elliptic in 2 or ovate tube broadly to corolla transversely lobes glabrate inside; 6 so base, long, sparsely the mm to above glabrate puberulent to just pilose outside, wide short sparsely, mm infundibular, corolla 1 pilose; ca. to flaring mi diameter, in mm 0.8–1 long, mm 1–1.8 section, cross in terete 0.2 the long, of mm portion free the 0.7 from hypanthium separating staminodes apex; the mi imtr oaewt 5 with 2.8 lobate diameter, cupular, in stigma mm glabrous; 3–4 long, diameter, in 0.5 disc mm disc; 2–2.3 circular a by subtended glabrous, hnmtr,bodyelpodt lbs,4 stripes globose, to orange-brown ellipsoid orange-yellow broadly darker mature, with with when darker orange-brown yellow both with or immature, it, green stripes when reaching medium stripes not mature, green but longitudinal and with apex, immature bicolored towards when fruit base apex; 2.5–4.5 from and at base stripes at diameter diameter in in mm 1 mm ca. apex, wider narrower a to a base from upwards flaring gradually 6.5 hairs, ped- breakaway section, cross visible; in readily sulcate and icel ground above maturing geocarpic, in ntemri;fne-iesimtcpoetos0.7 projections stigmatic 0.6 finger-like long, margin; mm the on tions litc (3.8 elliptic, (1 lost, bristle sometimes hyaline the with or at base, persistent than the eye, fruit near naked the compressed laterally on the apex, denser to granular, basally bloom aculei minutely tuberculate, thin glabrous, not a moder- aculeate, with pericarp pulverulent the thick, appearing with very apex, at to blunt ately diameter, in cm 5.8 ndaee;bite0.7 bristle diameter; in t osihl rnaea ae ral ct oots at obtuse to acute broadly 2 base, Figures at unwinged. truncate apex, slightly to emargin- sculpturing, ate surface evident without smooth brous, ...Iw:Gona h SANrhCnrlRegional Central North USDA the at Grown Iowa: Paratypes—U.S.A. S CHAEFER ie sln steopaque the as long as times 0.3–3 long, mm –1.5 –)4.86.2 (2007) 2 mln,pls ratos-tioewith antrorse-strigose or pilose long, mm 120 . mi imtr lbaeotie corolla outside; glabrate diameter, in mm –6.5 –11 . mi imtr iueypbrln;style puberulent; minutely diameter, in mm –0.6 . mi imtr hr papillate. short diameter, in mm –0.8 . maoeteoay ailfr,0.7 papilliform, ovary, the above mm –1.2 / fhpnhu refo h vr;free ovary; the from free hypanthium of –1/3 mi imtr lboswt tf aculei, stiff with glabrous diameter, in mm –9 × eaeflowers Female 0m,ots n urnt at mucronate and obtuse mm, (3–)6.510 × m 1.2 mm, (2.2–)2.43 mlong. mm –1 mln,3 long, mm –5 –4. . mwide, mm 0.4–0.6 long, mm –1 . mi di- in mm 1.3–1.7 long, mm –1 0c og ioewt non- with pilose long, cm –20 0m og 0.8 long, mm –)410 eielt;pdclslaein sulcate pedicel pedicellate; Seeds –1.3 igrlk projec- finger-like –7 mi diameter, in mm –4 . mtik gla- thick, mm –1.8 –4.5 × elws white, yellowish mln,3.9 long, cm –6 mm, (0.3–)0.71 . mlong, mm –0.6 × . mm 0.3–0.5 .zambianus C. Fruit . mm –1.5 . mm –2.5 .anguria C. –3mm –1.5 735 not –4 –1 – Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 736 F IG .2. ooye I505597, PI Holotype, zambianus . Cucumis YTMTCBOTANY SYSTEMATIC irehe,Cak&Risa572 Reitsma & Clark Widrlechner, (NA). Vlm 33 [Volume Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. enpeae yteoiia olcosi 1984. in international have collectors unsuccessful, might original that the an vouchers by herbarium after prepared any been selected locate to were conducted was types search These US). P, 569 oe.C aefoe.D aefoe,oee oso icadstamens. and disc stami- show and to opened stigma, flower, style, Male disc, D. flower. show Male to C. opened nodes. flower, Female B. flower. 94(I550)b .F teeadG lnoia aaia 9k N km 79 Samahina, at 2006, Mlongoti Mar 31 G. Zambia, and Attere 11°12Ј Jun F. Mwinilunga, 17 A. of collected by originally 505608) seeds (PI from 1984 Iowa, Ames, Station, Introduction 2006, S Apr km 2006, 4 63 (NA), Mar Kabuka, 31 at Zambia, Mlongoti 11°54Ј Province, G. Kamapanda, western and to Attere F. Mwinilunga A. of by 505597) (PI 1984 Jun 2008] b HJ,P 505597, PI JHKJ), (by F IG IC,2 p 2006, Apr 20 (ISC), 3 A .3. F IG –D. .4. uui zambianus Cucumis 568 rs ris nldn rs eto n aa iw I505608, PI view. basal a and section cross a including fruits, Fresh zambianus. Cucumis N) 0Ar2006, Apr 20 (NA), irehe,Cak&Risa567 Reitsma & Clark Widrlechner, irehe,Cak&Risa572 Reitsma & Clark Widrlechner, ,24°15Ј S, 571 IC,3 a 2006, May 31 (ISC), ,ee.110m otwsenProvince, Northwestern m, 1,180 elev. E, rw rmhltp n isotypes and holotype from Drawn . ,24°8Ј S, irehe,Cak&Risa566 Reitsma & Clark Widrlechner, 570 IREHE TA. UUI ZAMBIANUS CUCUMIS AL.: ET WIDRLECHNER N) rw tUD Plant USDA at grown (NA); 573 ,ee.130m North- m, 1,320 elev. E, B,IC ,M,NA, MO, K, ISC, (BR, IC,1 p 2006, Apr 11 (ISC), N) .Female A. (NA). ftecutyi hc twsdsoee,Zambia. discovered, was it which in country the of eaten. and peanuts, crushed and soda with mixed boiled, are as 505597 PI of name common r itntv ohfoitclyadpyigoial.They physiognomically. and floristically both distinctive are by dominated woodland area miombo an bezian through scattered widely farms crops and Moench, local of and collections fertility included Seed soil malnutrition. poor local reported extensive villages. (1986) isolated Kalumba and fields and farm re- Freund few with this low, in very production was agricultural gion exploration that this indicate from notes 1984) The (Attere plant. garden cultivated, a as Kalumba and (Freund re- there who communities 1986). ethnicity, Lunda Lunda of in mainly 25,000) settled Zaire, (perhaps North- and the refugees Angola collections, many from these received of time Province in the western addition, preceding yr In 20 boundaries. the international those White spans of which area northern the in phytochorion found is the It well. in ac- 5). nearby four Fig. in collected other circles The also (open 5). Province were Fig. Northwestern in 1) 15 circle was (Appendix 505608 (filled PI cessions Zaire and 14 of 5), collected Fig. west in was km star 505597 (filled PI Angola of apart; east collections km paratype 78 made and were holotype The System. Germplasm of collections no 1984), bianus (Attere Attere 1984 by Zambia in of Mlongoti parts and other from collected seed were cucurbit samples many on-farm Although from gardens. times 5), backyard six and (Fig. region storage the Zambia from of sampled was Province it Northwestern where the of corner ern Distribution Etymology omnNm n Use and Name Common ed o nwi hstxnocr nawl tt ronly or state wild a in occurs taxon this if know not do We zambianus Cucumis rmohrrgoswr upidt h ainlPlant National the to supplied were regions other from uubt maxima Cucurbita h pcfceihti omdfo h name the from formed is epithet specific —The e mays Zea t nw itiuini ntenorthwest- the in is distribution known —Its ‘ abza einlcnr fendemism, of centre regional Zambezian rbbyocr nAgl n ar as Zaire and Angola in occurs probably .(tee18) n eemd from made were and 1984), (Attere L. Wie18,18) hs woodlands These 1983). 1981, (White irehe,Cak&Risa573 Reitsma & Clark Widrlechner, — Duchesne, ‘Katanda h olcosgv h local the gave collectors The oigta t seeds its that noting ’, ogu bicolor Sorghum 18,1983) (1981, ’s . 5km –15 .zam- C. Zam- (L.) 737 –17 ’ Delivered by Publishing Technology to: Iowa State University IP: 129.186.176.91 on: Thu, 17 Jan 2013 08:28:31 Copyright (c) American Society for Plant Taxonomists. All rights reserved. xliigteuiiyo o oyncergnsin genes nuclear copy low of utility the exploiting of to bianus placement needed phylogenetic is the genes determine additional confidently employing species, related comprising the clade a within resolution of lack nnmu eiwr o hi epu ugsin nrveigthis reviewing in suggestions Clark, two helpful and G. Wiersema, their Lynn H. manuscript. for John Drs. and reviewers and Stevens, F. anonymous report, Peter exploration Robertson, R. original Kenneth assistance cultivation, the for plant Thormann supplying in Imke assistance in Ms. technical preparation, like voucher for would and Clark the pollination, We Lucinda by agencies. Ms. product cooperating thank any or to of Agriculture endorsement of commercial an Department of constitute Mention U.S. not funds. Iowa does of names State was brand and 1018, Act No. Hatch Project by Iowa, supported Ames, Station, Experiment Economics Home ewrhhl oepn neseii rsigexperiments crossing would include interspecific it to expand addition, to In worthwhile cultivation. be of outside populations for cur- of the extent determine rent to areas surrounding and Zambia evergreen. western almost to deciduous from semidecidu- vary typically can are but ous, They light, a crown. branches form flat-topped to lower spreading shallow, branches trunks, upper and slender erect, strongly short, of with species by appearance dominated always almost and Brachystegia tall m 10–20 are 738 aba ildsa:P 057 ildcrl:P 068 pncrls addi- circles: open 1. 505608; Appendix PI in circle: listed filled populations 505597; tional PI star: Filled Zambia. A F uueResearch Future IG CKNOWLEDGMENTS .Koncleto ie for sites collection Known 5. . mn t ognr.Tetidato scurrently is author third The congeners. its among .insignis C. et.(aaee,wihaedsiciei their in distinctive are which (Fabaceae), Benth. .zambianus C. hsjunlppro h oaArclueand Agriculture Iowa the of paper journal This . twudb aubet eunt north- to return to valuable be would —It , .dipsaceus C. utvto n s,adt search to and use, and cultivation uui zambianus Cucumis and , .anguria C. .zambianus C. nnorthwestern in ie the Given . YTMTCBOTANY SYSTEMATIC Cucumis .zam- C. and . ars .S 99 h eeto ne n h ecldcnitnyindex. consistency rescaled the and index retention The 1989. S. J. Farris, wnlnat yksy,Nrhetr Province. Northwestern Nyakaseya, to Mwinilunga Province. Northwestern Mwinilunga, of zambianus S km 67 mapanda, 2006. as identified previously ration esnti,J 95 ofdnelmt npyoeis napoc using approach an phylogenies: on limits Confidence 1985. J. Felsenstein, lr,R . .P irehe,C .Bok n .R etm.19.Cu- 1991. Reitsma. R. K. and Block, C. C. Widrlechner, P. M. L., R. Clark, 1984. F. A. Attere, wnlnat yksy,Nrhetr Province. Northwestern Nyakaseya, to Mwinilunga anus ikrd,J .J.1993. Jr. H. J. the Kirkbride, of evolution Molecular 1998. Hemleben. V. and King, K. J., Jobst, 13. 1978. Jeffrey. C. and P. Halliday, of Relationships 2007. Barber. C. J. and Thulin, M. G., in A. refugees Ghebretinsae, settled Spontaneously 1986. Kalumba. K. and J. P. Freund, eg oNaaea otwsenProvince. Northwestern Nyakaseya, to lenge anus imn,M .adH coeea 00 asa hrcesi sequence- in characters as Gaps 2000. Ochoterena. H. and P. M. Simmons, 2007. Ox- H. v2.0a11. Schaefer, editor, alignment evolu- Sequence the Se-Al: and 1996–2002. phyletics A. Quantitative Rambout, 1969. Farris. S. J. and G. A. Kluge, g,piiieclia/adae ymol,6 mfo unoft Gy- to off turn Province. from Southern km embe, 67 Syamyoolo, cultivar/landrace; primitive age, ht,F 1983. F. White, 1981. F. parsimony White, using analysis Phylogenetic PAUP*: 2002. L. D. Swofford, I505605 PI A PPENDIX Cladistics ubtgrpama h ot eta einlPatIntroduction Plant Regional Iowa. Central Ames, North Station, the at germplasm curbit for Board Resources. International Genetic Rome: Plant IBPGR/84/114. AGPG: Report FAO ora fBotany of Journal oeeisadEvolution Cucurbitaceae. and family logenetics rela- the phylogenetic of and species ITS2) among tionships and (ITS1 Spacers Transcribed Internal Man- Zambesiaca Flora London: Launert. Committee. E. aging ed. 4, vol. Zambesiaca, of phylogenetics molecular cumis unraveled: melons and cucumbers 299–312. 20: Zambia. Province, Northwestern bootstrap. the ae hlgntcanalyses. phylogenetic based data. DNA plastid and nuclear on based tion Myrmecosicyos Mukia, coelospermum, Oxford. of University U.K.: ford, Anurans. of tion (Cucurbitaceae) ai:UNESCO. Paris: Africa. of map vegetation UNESCO/AETFAT/UNSO Associ- Sinauer Sunderland: 4.0b10. ates. Version methods), other (*and amsoae rmtv utvrlnrc;Ktk,1 mfo Ike- from km 15 Katoka, cultivar/landrace; primitive storage, Farm amsoae rmtv utvrlnrc;Klmt,6 mNof N km 69 Kalimoto, cultivar/landrace; primitive storage, Farm amsoae rmtv utvrlnrc;Nhl,5 mNof N km 55 Nchila, cultivar/landrace; primitive storage, Farm n eae eea(eicsa,Cucurbitaceae). (Benincaseae, genera related and .Adtoa cesosfo 94IGIZminexplo- Zambian IPGRI 1984 from accessions Additional 1. .zambianus C. :417–419. 5: h eeaino fia ecitv eort copn the accompany to memoir descriptive A Africa: of vegetation The ai:UNESCO. Paris: 1:5,000,000. Scale Africa: of map Vegetation Cucumis on,NrhCrln:PrwyPublishers. Parkway Carolina: North Boone, . Evolution IBPGR/ Zambia. in mission collecting germplasm Crop 4 1256–1266. 94: ytmtcZoology Systematic isseai oorp fteGenus the of Monograph Biosystematic L HortScience Ccriaee utinclude must (Cucurbitaceae) akad rmtv utvrlnrc;Ka- cultivar/landrace; primitive Backyard, ITERATURE :204–219. 9: 9 783–791. 39: Cucumis ytmtcBiology Systematic 6 2,450 326, 26: h nentoa irto Review Migration International The p seautddrn h alof fall the during evaluated as sp. Cucumis C and , 8 1–32. 18: ITED I505610 PI recircumscrip- A Oreosyce: 7 in 462–470 Pp. L. –451. Blumea 9 369–381. 49: I505607 PI I505609 PI .melo C. uuel,Di- Cucumella, oeua Phy- Molecular Vlm 33 [Volume 2 165–177. 52: I505606 PI amstor- Farm .zambi- C. .zambi- C. American Cucumis Flora Cu- C.