TAXON: Annona hypoglauca Mart. SCORE: 2.0 RATING: Low Risk

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1 Taxo: Aoa hypoglauca Mart. Family: Aoaceae Commo Name(s): ahuaracatoco Syoym(s): Aoa tessmaii Diels aoilla chirimoya guaábaa sacha majagua de aparo sacha-aoa wild cherimoya wild soursop Assessor: Chuck Chimera Status: Assessor Approved Ed Date: 7 Apr 2016 WRA Score: 2.0 Desigatio: L Ratig: Low Risk Keywords: Tropical Tree, Uarmed, Edible Fruit, Fleshy-Fruit, Fish-Dispersed Optio Aswer 101 Is the species highly domesticated? y=-3, =0 102 Has the species become aturalized where grow? 103 Does the species have weedy races? 201 Species suited to tropical or subtropical climate(s) - If islad is primarily wet habitat, the substitute "wet tropical" for "tropical or subtropical" (0-low; 1-itermediate; 2-high) (See Appedix 2) 202 Quality of climate match data (0-low; 1-itermediate; 2-high) (See Appedix 2) High 203 Broad climate suitability (evirometal versatility) y=1, = Native or aturalized i regios with tropical or subtropical climates Does the species have a history of repeated itroductios outside its atural rage? y=1, =0 y=-2,?=-1, =0 301 Naturalized beyod ative rage y = 1*multiplier (see Appedix 2), = questio Garde/ameity/disturbace weed =0, y = 1*multiplier (see Appedix 2) 303 Agricultural/forestry/horticultural weed =0, y = 2*multiplier (see Appedix 2) 304 Evirometal weed =0, y = 2*multiplier (see Appedix 2) 305 Cogeeric weed =0, y = 1*multiplier (see Appedix 2) y 401 Produces spies, thors or burrs y=1, =0 402 Allelopathic 403 Parasitic y=1, =0 404 Upalatable to grazig aimals High y Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 1 of 14

2 Optio Aswer 405 Toxic to aimals y=1, =0 406 Host for recogized pests ad pathoges 407 Causes allergies or is otherwise toxic to humas y=1, =0 408 Creates a fire hazard i atural ecosystems y=1, =0 409 Is a shade tolerat plat at some stage of its life cycle 410 Tolerates a wide rage of soil coditios (or limestoe coditios if ot a volcaic islad) 411 Climbig or smotherig growth habit y=1, =0 412 Forms dese thickets y=1, =0 501 Aquatic y=5, =0 502 Grass y=1, =0 503 Nitroge fixig woody plat y=1, = Geophyte (herbaceous with udergroud storage orgas -- bulbs, corms, or tubers) Evidece of substatial reproductive failure i ative habitat y=1, =0 y=1, =0 602 Produces viable seed y=1, =-1 y 603 Hybridizes aturally 604 Self-compatible or apomictic 605 Requires specialist polliators 606 Reproductio by vegetative fragmetatio 607 Miimum geerative time (years) 701 Propagules likely to be dispersed uitetioally (plats growig i heavily trafficked areas) y=1, = Propagules dispersed itetioally by people y=1, =-1 y 703 Propagules likely to disperse as a produce cotamiat y=1, = Propagules adapted to wid dispersal y=1, = Propagules water dispersed y=1, =-1 y 706 Propagules bird dispersed y=1, = Propagules dispersed by other aimals (exterally) y=1, = Propagules survive passage through the gut y=1, =-1 y 801 Prolific seed productio (>1000/m2) 802 Evidece that a persistet propagule bak is formed (>1 yr) 803 Well cotrolled by herbicides 804 Tolerates, or beefits from, mutilatio, cultivatio, or fire 805 Effective atural eemies preset locally (e.g. itroduced biocotrol agets) Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 2 of 14

3 Supportig Data: 101 Is the species highly domesticated? Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, Part 2. Papilioaceae, Mimosaceae, Coaraceae, Aoaceae, Additios ad Correctios. Foudatio Va Phillips, O. (1993). The potetial for harvestig fruits i tropical raiforests: ew data from Amazoia Peru. Biodiversity & Coservatio, 2(1), Missouri Botaical Garde Flora of the Veezuela Guayaa. Aoaceae. [Accessed 7 Apr 2016] 102 Has the species become aturalized where grow? NA 103 Does the species have weedy races? NA 201 Species suited to tropical or subtropical climate(s) - If islad is primarily wet habitat, the substitute "wet tropical" for "tropical or subtropical" USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database]. High "Native: Souther America Brazil: Brazil - Acre, - Amazoas, - Para, - Amapa, - Rodoia, - Roraima Norther South America: Frech Guiaa; Guyaa; Suriame; Veezuela - Amazoas Wester South America: Bolivia - Bei, - La Paz, - Pado; Colombia; Ecuador - Napo, - Sucumbios; Peru - Amazoas, - Loreto, - Madre de Dios, - Ucayali" Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 3 of 14

4 202 Quality of climate match data High USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database] Broad climate suitability (evirometal versatility) Missouri Botaical Garde Flora of the Veezuela Guayaa. Aoaceae. [Accessed 7 Apr 2016] [Low elevatio tropics] "Seasoally flooded riparia forests, ca. 100 m; Amazoas (Sa Carlos de Río Negro). Colombia, Guyaa, Suriame, Frech Guiaa, Peru, Amazoia Brazil, Bolivia." 204 Native or aturalized i regios with tropical or subtropical climates USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database]. y "Native: Souther America Brazil: Brazil - Acre, - Amazoas, - Para, - Amapa, - Rodoia, - Roraima Norther South America: Frech Guiaa; Guyaa; Suriame; Veezuela - Amazoas Wester South America: Bolivia - Bei, - La Paz, - Pado; Colombia; Ecuador - Napo, - Sucumbios; Peru - Amazoas, - Loreto, - Madre de Dios, - Ucayali" 205 Does the species have a history of repeated itroductios outside its atural rage? Trade Wids Fruit Tropical Fruit. "A exceedigly rare Aoa (or sometimes Raimodia) from the highlad raiforests of South America." of widespread cultivatio outside ative rage Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 4 of 14

5 301 Naturalized beyod ative rage Radall, R.P A Global Compedium of Weeds. 2d Editio. Departmet of Agriculture ad Food, Wester Australia Wager, W.L., Herbst, D.R.& Lorece, D.H Flora of the Hawaiia Islads. Smithsoia Istitutio, Washigto, D.C. ra/idex.htm. 302 Garde/ameity/disturbace weed Radall, R.P A Global Compedium of Weeds. 2d Editio. Departmet of Agriculture ad Food, Wester Australia 303 Agricultural/forestry/horticultural weed Radall, R.P A Global Compedium of Weeds. 2d Editio. Departmet of Agriculture ad Food, Wester Australia 304 Evirometal weed Radall, R.P A Global Compedium of Weeds. 2d Editio. Departmet of Agriculture ad Food, Wester Australia 305 Cogeeric weed y Weber, E Ivasive Plat Species of the World. A Referece Guide to Evirometal Weeds. CABI Publishig, Walligford, UK "Aoa glabra is a ivasive tree/shrub that forms dese thickets ad shades out ative shrubs ad trees by prevetig their establishmet ad growth. Species richess is reduced i stads of this tree/shrub." Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 5 of 14

6 401 Produces spies, thors or burrs [] "A shrub up to 10 metres high; youg brachlets ferrugious-tometose. Leaves o 7-11 mm log petioles. rather Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, membraaceous. smooth ad fially glabrous o upper side. whiteglaucous ad covered with small. lustrous. appressed hairs o the Part 2. Papilioaceae, Mimosaceae, Coaraceae, Aoaceae, Additios ad Correctios. Foudatio Va uder side. elliptic. ovate or oblog. rotudate or shortly acute at the base. rather log ad mostly gradually atteuate at the apex cm log ad cm broad." 402 Allelopathic Ukow 403 Parasitic Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, Part 2. Papilioaceae, Mimosaceae, Coaraceae, "A shrub up to 10 metres high" [Aoaceae. ] Aoaceae, Additios ad Correctios. Foudatio Va 404 Upalatable to grazig aimals Jaick, J.& Paull, R.E The Ecyclopedia of Fruit & Nuts. CABI Publishig, Walligford, UK [Fruit palatable to fish & humas. Palatability of foliage ukow] "The yellow fruits fall ito the water immediately upo ripeig ad float at the surface where they are eate by the fish, which swallow the etire seed"... "Because of its fie flavor, fruits of Aoa hypoglauca are also appreciated by ma" [Ukow. Related Aoa is palatable to aimals] "Aoa seegalesis wild soursop" "Youg plats do ot compete well with weeds ad eed to be protected from fire ad browsig aimals." 405 Toxic to aimals Quattrocchi, U CRC World Dictioary of Medicial ad Poisoous Plats: Commo Names, Scietific Names, Epoyms, Syoyms, ad Etymology. CRC Press, Boca Rato, FL Wagstaff, D.J Iteratioal poisoous plats checklist: a evidece-based referece. CRC Press, Boca Rato, FL 406 Host for recogized pests ad pathoges Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 6 of 14

7 Peña, J. E., Nadel, H., Barbosa-Pereira, M., & Smith, D. (2002). Polliators ad Pests of Aoa Species. Tropical Fruit Pests ad Polliators: Biology, Ecoomic Importace, Natural Eemies, ad Cotrol, 197. CAB Iteratioal, Walligford, UK [Geeric descriptio of pests. Specifics for A. hypoglauca ukow] "I the Neotropics 296 species of arthropods are recorded as associated with Aoa species. The families most frequetly observed o Aoa species are Coccidae (Homoptera), Noctuidae, Oecophoridae (Lepidoptera), ad Eurytomidae (Hymeoptera)." 407 Causes allergies or is otherwise toxic to humas Quattrocchi, U CRC World Dictioary of Medicial ad Poisoous Plats: Commo Names, Scietific Names, Epoyms, Syoyms, ad Etymology. CRC Press, Boca Rato, FL Wagstaff, D.J Iteratioal poisoous plats checklist: a evidece-based referece. CRC Press, Boca Rato, FL [] "Because of its fie flavor, fruits of Aoa hypoglauca are also appreciated by ma. The soft flesh of the fruit cotais starch." 408 Creates a fire hazard i atural ecosystems Missouri Botaical Garde Flora of the Veezuela Guayaa. Aoaceae. [Accessed 7 Apr 2016] [. Does ot occur i fire proe habitats] "Seasoally flooded riparia forests, ca. 100 m; Amazoas (Sa Carlos de Río Negro). Colombia, Guyaa, Suriame, Frech Guiaa, Peru, Amazoia Brazil, Bolivia." 409 Is a shade tolerat plat at some stage of its life cycle [Shade tolerace ukow] "Aoa hypoglauca occurs i the iudated forests (varzeas) of Suriam, ad Brazilia ad Bolivia Amazo..." 410 Tolerates a wide rage of soil coditios (or limestoe coditios if ot a volcaic islad) [Soil types ukow] "Aoa hypoglauca occurs i the iudated forests (varzeas) of Suriam, ad Brazilia ad Bolivia Amazoia..." 411 Climbig or smotherig growth habit Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 7 of 14

8 Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, Part 2. Papilioaceae, Mimosaceae, Coaraceae, "A shrub up to 10 metres high; youg brachlets ferrugioustometose." Aoaceae, Additios ad Correctios. Foudatio Va 412 Forms dese thickets Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, Part 2. Papilioaceae, Mimosaceae, Coaraceae, [] "Distributio: Suriame ad Amazo basi to Bolivia" Aoaceae, Additios ad Correctios. Foudatio Va 501 Aquatic [Not aquatic, but occurs i iudated forests] "Aoa hypoglauca occurs i the iudated forests (varzeas) of Suriam, ad Brazilia ad Bolivia Amazoia (Fries 1931:250)..." 502 Grass USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database]. "Family: Aoaceae" 503 Nitroge fixig woody plat USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database]. "Family: Aoaceae" 504 Geophyte (herbaceous with udergroud storage orgas -- bulbs, corms, or tubers) Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, Part 2. Papilioaceae, Mimosaceae, Coaraceae, "A shrub up to 10 metres high..." Aoaceae, Additios ad Correctios. Foudatio Va 601 Evidece of substatial reproductive failure i ative habitat Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 8 of 14

9 USDA, ARS, Germplasm Resources Iformatio Network, Natioal Plat Germplasm System [Olie Database]. [. wide distributio] "Native: Souther America Brazil: Brazil - Acre, - Amazoas, - Para, - Amapa, - Rodoia, - Roraima Norther South America: Frech Guiaa; Guyaa; Suriame; Veezuela - Amazoas Wester South America: Bolivia - Bei, - La Paz, - Pado; Colombia; Ecuador - Napo, - Sucumbios; Peru - Amazoas, - Loreto, - Madre de Dios, - Ucayali" 602 Produces viable seed y "I 31 aalysed species, percet germiatio after 7 weeks varied Paroli, P., Ferreira, L. V., & Juk, W. J. (2003). Germiatiobetwee 5 ad 100% i the seeds i well watered soil, with a characteristics ad establishmet of trees from cetral average of 50%."... "The fastest germiatio with the give Amazoia flood plais. Tropical Ecology, 44(2), coditios occurred i Salix humboldtiaa (3 days), the slowest i Aoa hypoglauca ad Hevea spruceaa (49 days)." 603 Hybridizes aturally Pito, A.C. de Q., Cordeiro, M.C.R., Adrade, S.R.M. de, Ferreira, F.R., Filgueiras, H.A.C., Alves, R.E. & Kipara, D.I Aoa species. Iteratioal Cetre for Uderutilised Crops, Uiversity of Southampto, Southampto, UK [Ukow. Hybridizatio possible i geus] "This slight variatio i chromosome umber may explai the ease or difficulty of iterspecific hybridisatio ad graftig, ad warrats further work to determie if itra specific variatio also exists. Some related species, e.g., A. glabra, are kow to be tetraploid (Kessler, 1993, cited by Scheldema, 2002). Geerally, crosspolliatio betwee aoas is coducted primarily to determie compatibility for icreasig fruit set (Nakasoe ad Paull, 1998) ad occasioally for ew hybrid developmet." Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 9 of 14

10 604 Self-compatible or apomictic Peña, J. E., Nadel, H., Barbosa-Pereira, M., & Smith, D. (2002). Polliators ad Pests of Aoa Species. Tropical Fruit Pests ad Polliators: Biology, Ecoomic Importace, Natural Eemies, ad Cotrol, 197. CAB Iteratioal, Walligford, UK [Possibly No. Geeric descriptio] "Aoaceous flowers are protogyously dichogamous, opeig as females with receptive stigmas ad closed athers, ad later losig stigmal receptivity as the flowers tur ito polle sheddig males (Gottsberger, 1970). This evolutioary adaptatio prevets depositio of polle o to the stigmas i the same flower, ad is oe of may techiques that plats employ to avoid self-fertilizatio. Prevetio of the trasfer of polle betwee differet flowers o the same plat is aother techique used by may Aoaceae, ad is achieved through sychroizatio of flowerig, where, at ay time, ope flowers o oe plat are fuctioally of oly oe sex. These temporal floral traits, commoly ehaced with iheret icompatibility betwee polle ad ovule from the same plat, reder most aoaceous species uable to self-polliate." 605 Requires specialist polliators va Dulme, A. (2001). Polliatio ad Pheology of Flowers i the Caopy of Two Cotrastig Rai Forest Types i Amazoia, Colombia. Plat Ecology, 153(1/2), Peña, J. E., Nadel, H., Barbosa-Pereira, M., & Smith, D. (2002). Polliators ad Pests of Aoa Species. Tropical Fruit Pests ad Polliators: Biology, Ecoomic Importace, Natural Eemies, ad Cotrol, 197. CAB Iteratioal, Walligford, UK "Appedix I Life form, polliators ad sexual system of the plats foud i the caopy of a uplad rai forest ad a seasoally iudated rai forest i Amazoia, Colombia" [Aoa hypoglauca - Polliator = beetles] [Possibly polliator limited] "Iadequate polliatio is implicated as a major factor limitig productio of commercial Aoa fruits i may locatios (Gazit et al., 1982). This is attributed, i part, to the temporal separatio of female ad male fuctio withi the flower, which limits its potetial to self-polliate without exteral factors. The commoest problem is lack of polliators. This is a direct result of the expasio of platatios ito regios outside the ative rage of the plats ad their polliators, ad may be due eve to the uatural coditios imposed by cultivatio, regardless of locality."... "The majority of Aoaceae are polliated by beetles, although some are polliated by thrips (e.g. Momose et al., 1998), true bugs (Farre et al., 1997), ad eve cockroaches (Nagamitsu ad Ioue, 1997)." 606 Reproductio by vegetative fragmetatio Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, "A shrub up to 10 metres high" [Ukow. A tree reported to Part 2. Papilioaceae, Mimosaceae, Coaraceae, reproduce by seed, but ukow whether also able to reproduce Aoaceae, Additios ad Correctios. Foudatio Va vegetatively] Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 10 of 14

11 607 Miimum geerative time (years) Ukow 701 Propagules likely to be dispersed uitetioally (plats growig i heavily trafficked areas) Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, [Fish-dispersed. Fruit & seeds lack meas of exteral attachmet] Part 2. Papilioaceae, Mimosaceae, Coaraceae, "Fruit elogate-ellipsoid. fulvous-tometellous. 5 cm log ad 2.5 cm Aoaceae, Additios ad Correctios. Foudatio Va across; areoles elogate, 2-3 mm broad. taperig ito short. more or less up curved Ad ofte evetually disappearig cusps." 702 Propagules dispersed itetioally by people y Trade Wids Fruit Tropical Fruit. [Seeds sold olie] "1 seed per pack. A exceedigly rare Aoa (or sometimes Raimodia) from the highlad raiforests of South America. Bears smallish, yellow ripeig fruits that are likely edible, much like its may relatives." 703 Propagules likely to disperse as a produce cotamiat [. Ulikely give large fruit & seed size] "The black seed has a whitish, slimy arillus-like out- growth (arilloid) of cartilagious cosistecy. Seed ad arilloid together are 1.3 cm log; the arilloid aloe is 0.3 x 0.5 cm."... "The yellow fruits fall ito the water immediately upo ripeig ad float at the surface where they are eate by the fish, which swallow the etire seed." 704 Propagules adapted to wid dispersal Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, [. Fleshy-fruited] "Fruit elogate-ellipsoid. fulvoustometellous. 5 cm log ad 2.5 cm across; areoles elogate, 2-3 Part 2. Papilioaceae, Mimosaceae, Coaraceae, Aoaceae, Additios ad Correctios. Foudatio Va mm broad. taperig ito short. more or less up-curved Ad ofte evetually disappearig cusps." 705 Propagules water dispersed y [Water & fish dispersed] "Aoa hypoglauca fruits ripe as the fish become more commo. The yellow fruits fall ito the water immediately upo ripeig ad float at the surface where they are eate by the fish, which swallow the etire seed." 706 Propagules bird dispersed Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 11 of 14

12 [Fish-dispersed] "The yellow fruits fall ito the water immediately upo ripeig ad float at the surface where they are eate by the fish, which swallow the etire seed." 707 Propagules dispersed by other aimals (exterally) Lajouw, J. & Stoffers, A. L Flora of Suriame. Vol. II, [Fish-dispersed. No meas of exteral attachmet] "Fruit elogateellipsoid. fulvous-tometellous. 5 cm log ad 2.5 cm across; areoles Part 2. Papilioaceae, Mimosaceae, Coaraceae, Aoaceae, Additios ad Correctios. Foudatio Va elogate, 2-3 mm broad. taperig ito short. more or less up-curved ad ofte evetually disappearig cusps." 708 Propagules survive passage through the gut y Maia, L. A., Maia, S., & Paroli, P. (2005). Seedlig morphology of o-pioeer trees i Cetral Amazoia várzea floodplai forests. Ecotropica, 11(1-2), 1-8 "I Ziburski s study, A. hypoglauca germiated oly after submergece or passage through a fish digestive system, whereas i our study this species germiated i 60% of cases." [Dispersed by fish & water] "Aoa hypoglauca fruits ripe as the fish become more commo. The yellow fruits fall ito the water immediately upo ripeig ad float at the surface where they are eate by the fish, which swallow the etire seed. Fishes which are alleged to eat the fruits ad seeds of A. hypoglauca, ad i the stomach of which fisherme frequetly fid etire ubroke seeds, are: tambaqui, jatuaraa, pacu braco, pacu, ad sardiha. Because of its fie flavor, fruits of Aoa hypoglauca are also appreciated by ma. The soft flesh of the fruit cotais starch. We sometimes foud beetle larvae iside the ripeig fruits, which are presumably also eate by the fish." 801 Prolific seed productio (>1000/m2) Ukow 802 Evidece that a persistet propagule bak is formed (>1 yr) Trade Wids Fruit Tropical Fruit. Royal Botaic Gardes Kew. (2016) Seed Iformatio Database (SID). Versio "Seeds are fresh, caot be stored ad should be plated o receipt." "Storage Behaviour: No data available for species. Of 8 kow taxa of geus Aoa, % Orthodox(p/?)" 803 Well cotrolled by herbicides Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 12 of 14

13 Ukow. No iformatio o herbicide efficacy or chemical cotrol of this species 804 Tolerates, or beefits from, mutilatio, cultivatio, or fire Ukow 805 Effective atural eemies preset locally (e.g. itroduced biocotrol agets) Ukow Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 13 of 14

14 Summary of Risk Traits: High Risk / Udesirable Traits Thrives i tropical climates Other Aoa species are ivasive Reproduces by seeds Seeds dispersed by fish, water & itetioally by people Limited ecological iformatio may reduce accuracy of risk predictio Low Risk Traits No reports of ivasiveess or aturalizatio, but o evidece of widespread itroductio outside ative rage Uarmed (o spies, thors or burrs) No-toxic Edible fruit Relatively large fruit & seeds may miimize risk of accidetal dispersal Secod Screeig Results for Tree/tree-like shrubs (A) Shade tolerat or kow to form dese stads?> No. Not kow to form dese stads. Shade tolerace ukow (B) Bird or clearly wid-dispersed?> No Outcome = Accept (Low Risk) Creatio Date: 7 Apr 2016 (Aoa hypoglauca Mart.) Page 14 of 14

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