"Terra Firme" Forests in Peruvian Amazonia

1’ It IN C 11’ ES [VOL. 35

Prirtriprs, 35(1), 1991, pp. 22-26 The Palm Communities of Two “Terra Firme” Forests in Peruvian Amazonia

FRANCIS%hHN AND KEMUERMEJIA ORSlnlll, /Ipurtudo postal 18-1209, Limu, Peru, and IIAP. Apartudo postal 784, Iquitos, Peru

AUSTIIACT (4q055‘S; 73O44O‘W). The first site, called LLFerrocako,” is about 6 lcm north of the Two pnlm surveys cnrried out in “terra firiiio” forests of ilio lower Ucnynli Ilivcr volley in Peruvian field staiion of the Research Institute for Aniuzoniii rcprt very l~iglidiversities \vitIl, respec. Peruvian Amazonia (IIAP); the seocnd, tivcly, 29 slicciics in i(, gcncro on 0.71 ha a:d 34 called “Copal,” is at Icm 15 on the only sliccics in 21 genrru on 0.5 ha. 111 both cnses, inost road. Both sites belong to the same geo- sliccics iiro sinnll, understory p[tlms, ond niai e than graphic unit at an elevation of about 160 99% of tlic comniuiiity is utidcr 10 m in height. Severn1 grncro und Rpccics inventoried, rvhich nre not m above sea level. frequcnt in I’cruviun Amnzonia nt low elevations nnd Climate is humid tropical, with an aver- rutlicr cniiirwn on the Atitlt~nn piedmont arid on age annual rainfall of 2.9 m and average ~il~Jiillillili!lRlqwq, p~intout tho ilifluencc of IIICsuban. annual temperature of 26” C. , dciiri Iinlni floru on tlic wcstclri lowlands of the Am. milinsin. Soils of both sites correspond to acrisol in the FAO-UNESCO (197 1) classification “Terra firme” forests, which cover the system. They are yellow and clayey in their major part of the Amazon bc.sin, are con- upper part; brownish to dark brown at sidered a species rich ecosystem (ßlack et greater depth; the upper 4 meters are never al. 1950, Boom 1986, Gentry 1982, Gen- waterlogged; the drainage is vertical. At try and Dodson 1987, Pires et al. 1953, the Copal site, the soil differs slightly, hav- Prance et al. 1976). Previous reports from ing a silty layer in the upper horizon at French Guiana (Granville 1978, Sist 1985) the lowest part of the slope, and a sandy and from Brazil (Kahn 1986, Kahn and clay texture, at some places on the plateau Castro 1985) presented data on species at the margin of the area surveyed, cor- richness and density of palms in eastern responding to a transition zone into an area and central Ammonia, pointing out par- of gleyic podzol which extends about 50 ticularly the high diversity of the palm m beyond. community in a “terra firme” forest of the central basin; but there were no quanti- Methods tative data from western Amazonia. Trying Palm surveys. All palms were counted to fill the gap, we prcsent here two palm on contiguous 0.01-ha plots. For multi- survcys carried out in “terra firme” forests stemmed species, all axes were counted. in Peruvian Amazonia, including data on Total areas of 0.71 ha and 0.5 ha were species richness, density, life forms and surveyed respectively at the Ferrocaño and vcrtical distribution of the palm commu- Copal sites. Density of each species is given according to height classes (under 1 m, 1 to 10 m, above 10 m), and lumped on the whole area at each site. ,..9 -o, Both forests are located in the loiver Palm collection and identijication. Ad o “x, Ca Ucayali River valley, near Jenaro Herrera species were collected. Some of them could lY911 ‘ KAIIN AND MEJIA: PALM COMMUNITIES 23 not be identified due to the lack of com- M 1999, Y”‘):Maximiliana maripa (Co- parative material in the herbaria (hP’rhZ, rrêa de Serra) Drude (Mejia 780, USM); ’

BH, I(, NY, USM). J. Dransfield contrib- + Oenocarpus balickii Kahn (K& M 1723, uted to the identification of many palms. NY); O.^mapora Karsten (K & LM 1727, Several names of Bactris are provisional; NY); Orbignya polysticha Burret (K & this genus badly needs a new taxonomic A1 1724$,NY); Phytelephas macrocarpa treatment. Voucher numbers and herbaria Ruiz et Pavon (K & M 1726, AAU); Pho- where plants are deposited are given in the Zidostachys synanthera (Martius) H. E. following list: Aiphanes ulei Burret (Kahn Moore (IC & M 1890, BH); Scheelea bas- 6% Mejia 1916, K); As.irocaryum cham- sleriana Burret (IC & M 1794*, USM); bira Burret (IC & M 1768, USM); A. Socratea exorrhiza (Martius) Wendland macrocalyx Burret (I<& M 1782, USM); (K & M 1702, NY); S. salazarii H. E. Bactris acan thocarpoides Barbosa Moore (K & M. 2013, BH); Wettinia Rodrigues (IC 6% M 1760, IC); B. acan- aupsta Poeppig et Endlicher (K & M thospatha Trail ex Drude (IC & M 1737, 1767, USM). K); B. bijda Martius (K & M 1789, K); B. humilis (Wallace) Trail (I< & M 1758, Results IC); B. cf. hylophila Spruce (I(& M 1798, K); B. mitis Martius (K & M 1763, K); At the Ferrocaño site, 29 species in 16 B. piranga Trail (K & M 1761, K); R. genera were encountered on 0.71 ha (Table simplicfrons Martius (IC & M 1781, 1). The genus Bactris is the most diver- USM); B. sphaerocarpa Trail (K di &I sified, with 10 species. Palm diversity var- 1813, K); B. sp. aff. B. mitis Martius (K ies from 7 to 144 species and 5 to 1O genera & M 1799, IC); B. sp. 1 (I<& M 1835, per 0.01 -ha plot. Palm density is very high IC); B. sp. 2 (K & 111 17419, K); B. SP.’~. (986 palms per 0.1 ha). While species (I< & &I 1905, USM); B. sp. 4 (K & M richness and density are both remarkably 1904, 10; Charnaedorea integrifolia high, the palm community is largely dom- (Trail) Dammer (IC & M 1790, K); Chel- inated by two small, multi-stemmed, yocarpus repens Kahn et Mejia (K & M understory species, Lepidocaryum tess- 1974, NY); Desmoncus cf. leptospadix marinii and Bactris sphaerocarpa, which Martius (K & M 1921, K); Euterpe pre- represent, respectively, 54.3% and 17.7% catoria Martius (IC & M 1778, USM); of the community. Geonoma acaulis Martius (K & M 2001, At the Copal site, 34 species in 21 gen- USM); G. camana Trail (I< & M 1765, era were encountered on 0.5 ha surveyed K); G. leptospadix Trail (K & M 1709, (Table 2). Geonoma with 8 species is the K); G. oligoclona Trail (K & M 1910, most diverse genus. Palm diversity varies 10; G. piscicauda Dammer (K.& M 181 O, from 6 to 18 species and 5 to 13 genera K); G. poeppigiana Martius (IC& M2020, per 0.01-ha plot. Of the 50 plots surveyed, BH); G. pycnostachys Martius (I( & M 27 present 12 or more species, and 31 1698, K); G. spixiana Martius (K & M have 9 or more genera. Palm density is 1808, IC); G. cf. tamandua Trail (K & M slightly lower than in the former site (768 1888, BII); Hyospathe elegans Martius palms per O. 1 ha). Lepidocaryum tess- (IC & M 1917, K); H. cf. weberbaueri manni¿ represents only 7.4% of the com- Dammer ex Burret (K & M 1815, K); munity. Astrocaryum macrocalyx and Zriartnn ddtoìdea Ruiz et Pavon (K & M Phytelcphas macrocarpa were only found 1766, USM); Zriartella stenocarpa Bur- on the lowest part of the slope where the ret (K & M 1718, NY); Jessenia bataua soil differs in having a silty layer in the (Martius) Burret (IC & M 1792, IJSM); upper horizon; both species are frequent Lepidocaryum tessmannii Burret (K & on alluvial soils. The four adults of Jes- Ta,ble I. Palm species richness and denuity at FerrocaZo site on 0.71 ha. S. single- stemmed; M, multi-stcmniecl; Acs, acaulescent with small leaves; Acl, acadescent with large leaves; L, lianescent; P, procumbent; LU, lower understory (adult height less than G m); UU, upper understory (adult height 6-10 m); AR, arborescent (adult height above 10 m); N, total number of palms.

Height Classes I~ulruSpecies . 10m N Life Form ds/rocnryim clin ni Oirn O 1 O 1 S.AR Un dris n en ri I h oca rpidcs 2 2 O 4 M.LU 13nclri.T fiiitii;/is O 1 O 1 M.LU I3nr1ri.v cf. hylophiln 3 11 O 14 M:LU BurIris niilis 2 10 O 12 M.LU f~flf"/l'iS /Jirf7ilfifi O 1 O 1 M.LU Bnr/ris siniplirifions 1 2 O 3 M.LU Bo cl ris sph (i r roc n rpi 339 898 O 1,237 M.LU Ilnrlris sp. off. n. nii/is ,2 3 O 5 M.LU ßnr/ris sp. 1 9 14 O 23 M.LU 13nclris sp. 2 12 101 O 113 M.LU Chrlyocnr~~i~srepens 18 lß8 O 206 S.P.LU Drsrnoiicrrs d. Irplospndir 1 1 O 2 M.L.UU Eirtrrpe prrcaforia 28 8 O 36 S.AR Grononin acnirlis 17 53 O 70 Acs.LU Grorionin 1rplospntii.v 15 22 O 37 M.LU Gtonorrrn pisciccmdn 70 102 O 178 M.LU Grononin porppigiorin 22 80 O 102 S.LU Ceononin spixinnn 57 191 O 248 M.LU Ifybspn/he cf. roebcrbnirrri 5 15 O 20 M.LU Irinrtclla strnocnrpn I20 67 O 187 M.LU Jcssrnh 6n1nim 151 195 O 34.6 S.AR Lrpidornryirm lr~siricirinii 4,55 3,346 O 3,801 M.LU il hiniìlin ti íi ni clrip n O 1 O 1. S.AR Orriocarpris bnliclzii 6 3 4 13 S.AR Clrhipiyn piilys/icfin 8 182 O 190 Acl.LU Pholidos/nrliys synnri/hrrn 27 117 O 144 S.LU Sorrnfen rsorrhizn 2 O 1 3 S.AR IVr//inin augrisla -1 -5 -O -6 M.UU 1.379 5,620 5 7,004 senia bataua were localed at the limit with caño (19.7%) and high at Copal (62.4%). a gleyic podzol, on which this species forms In the first site, the low value, due to the a dense population beyond the area sur- low density of most species, is accentuated veyed. by the high density reached by Lepido- At both sites, most OF the species are caryum tessmannii and Bactris sphaero- small, understory palms (23/29 at Ferro- carpa between 1 and 10 m. In the second caño; 27/34, at Copal), and most of the site, the high value is due to the high palm commurrity is distributed under 10 density of seedlings of two arborescent spe- m in height (99.9% at Ferrocaíío; 99.3% cies, Iriartea deltoidea and Jessenia at Copal). The density of large palms (above bataua, and of juveniles of three small, 10 m ' - kight) is higher ai Copal, 25 on understory species, Hyospatlze elegans, 0.5 ha, than at Ferrocaño, which had five Chamaedorea integriJolia, and Geonoma individuals on 0.71 ha. The percentage of piscicauda, which are particularly abun- palms nndcr 1 m in height is low at Ferro- dant in the clearings.

It '4 19911 1 KAHN AND MEJIA: PALM COMMUNITIES 25

Table 2. Palm species richness and density at Copal site on 0.5 ha. (Legend: see , '. Table 1).

I Height Classes Palm Species <1 m 1-10 m ,> 10 m N Life Form Aiphanes dei 2' 5 O 7 Acs.LU Astrocaryum chambira 14 21 2 37 S.AR Astrocarywm nzacrocalyx 31 66 O 97 s.uu Bactris acanthospatha 3 10 O 13 M.LU Bactris bijda 7 13 O 20 M.LU . Bactris piranga 2 2 O 4 M.LU Bactris sp. 3 a 17 O 25 M.LU Bactris sp. 4 1 7 O 8 M.LU Chamaedorea integrijolia 105 103 O zoa S.LU Chelyocarpus repens 7 14 O 21 S.P.LU Desmoncm cf. leptospadix 4 24 O 28 M.L.UU Euterpe precatoria 70 21 O 91 S.AR Geonoma acaulis 46 53 O 99 Acs.LU Geonoma camana 21 22 O 43 S.LU \ Geonoma oligoclona 33 '9 O 42 M.LU Geonoma piscicauda 94 65 O 159 M.LU Geonoma poeppigiana 32 42 O 74 S.LU Geonoma pycnostachys 30 47 O 77 M.LU Geonoma spixiana 32 52 O 84 M.LU Geonoma cf. tamandua 4 9 O 13 Acs.LU Hyospathe elegans 415 ' 221 O 636 M.LU Iriartea deltoidea 919 9 19 947 S.AR Iriartella stenocarpa 28 . 17 O 45 M.LU Jessenia bataua 260 , 161 4 425 S.AR Lepidocarycm tessrnannii 43 240 O 283 M,LU Maximiliana maripa 34 27 O 61 S.AR Oenocarpus mapora O 5 O 5 M.UU Orbignya polysticha O 1 O 1 Acl.LU , Pholidostachys synanthera 31 51 O a2 S.LU Phytelephas macrocarpa O 18 O ia M.LU Scheelea bassleriana 18 13 O 31 S.AR Socratea exorrhiza 14 8 O 22 S.AR Socratea salazarii 81 36 o. 117 SUU Wettinia augusta 7 -a -O -15 M.UU 2,396 1.417 25 3.838

at Copal site brings a new record: 34 spe- Discussion cies in 21 genera on 0.5 ha-with up to Both palm communities are character- 18 species on 0.01 ha, i.e., more species ized by a very high diversity. Data from than on 3.84 ha surveyed in the lower . the Ferrocaño site were previously com- Tocantins River valley. Three genera found pared with those from central and eastern in the area surveyed at the Copal site, Amazonian forests (Kahn et al. 1988): 29 Aiphanes, Chamaedorea, and Iriartea, species in 16 genera on 0.71 ha in the are not frequent at low elevations (less than lower Ucayali River valley; 26 species in 200 m above sea level) in Peruvian Ama- 9 genera on 0.72 ha in the lower Rio Negro zonia, while they are rather common in valley, near Manaus; and 12 species in 8 the forests of the Andean piedmont and genera on 3.844 ha in the lower Tocantins on mountain slopes. A fourth genus, Dic- valley, eastern Amazonia. The inventory tyocaryum (M & K 2010, USM), not i *,, . 81 26 PRINCIPES [VOL. 35

inventoried in the plois, occurs in the same FAO-UNESCO. 1971. Soil map of the world. Vol. forest near the area surveyed. This genus IV, South America. UNESCO, Paris. GENTRY,A. H. 1982. Patterns of neotropical plant was previously known only from higher species diversity. Evol. Biol. 15: 1-84. elevations (1,800-2,200 m) in Peru (Gen- . 1986. Notes on Peruvian palms. Ann. try 1986). The occurrence of these four Missouri Bot. Gard. 73: 158-165. genera, as well as that of certain species . AND C. H. DODSON. 1987. Contribution (Socmtca salazarli and Wcttinia a,ugusta) of nontrees to species richness of tropical rain forest. Biotropica 19: 149-156. which are frequcnt on the Andean pied- GRANVILLE, J. J. DE. 1978. Recherche sur la flore mont, paints out the influence of the suban- et la v6g6tation guyanaises. Thèse Doctorat dean palm ílora on the western lowlands d’Etat, Univ. Montpellier. 272 p. of the Amazon basin. As a result, palm KAHN,F. 1986. Les palmiers des forêts txepicales humides du bas Tocantins (Amazonie brksi. diversity is high in the Peruvian Amazonia. lienne). Rev. Ecol. (Terre et Vie) 4.1: 3-13. AND A. DE CASTRO. 1985. The palm com- Acknowledgment munity in a forest of central Amazonia, Brazil. Biotropica 17: 210-216. This study was supportcd by the inter! ,I<. MEJIA,AND A. DE CASTRO. 1988. ’ Spe- national agreement ORSTOM, France/ cies richness and density of palms in tetra firme IIAP, Peru. Soil descriptions were made forests of Amazonia. Biotropica 20: 266-269. by L. Veillon. We are indebted to I<. and PIRES, J. M., TH. DODZHANSKY, AND G. A. BLACK. 1953. An estimate of the number of species of 1-1. Clark for their helpful assistance on the trecs in an Amazoninn forest community. Bot. English manuscript. GU. 114: 467477. PRANCE, G. T.,w. A. RODRIGUEZ, AND M. F. DA LITERATUIIECITED SILVA.1976. Inventario florestal de um hect- are de mata de terra firme, km 30 da estrada BLACK,G. A., TII. DOIIZllANSKY, AND c. PAVAN. Manaus-Itacoatiara. Acta Amazonica 6(1): 9- 1950. Some attempts to estimate species diver- 35. shy and population density of trees in Amazonian SIST, P. 1985. Rég6n6ration et dynamique des forests. Bot. Gaz. 111: 413-425. populations de quelques espèces de palmiers en DOOM, B. M, 1986. A forest inventory ín Ama- forêt guyanaise. DEA Univ. Paris VI. 60 p. zonian Bolivia. Diotropica 18: 287-294.

Prirtcipcs, 35(1), 1991, p. 26

PALM LITERATURE MINTER, SUE, with contributions from CHRIS JONES, PETER MORRIS, and PETER RIDDINGTON.. 1990. The Greatest Glasshouse: the Rainforests Recreated. London: IINISO. L25.00. This uttractivc book is divided into two parts. The first part “Greenery among the iron: thc Palm Ilouse and its c011cctions~’has ten chapters. The first three discuss the origin of the Palm I-Iousc, thc dcvclopincnt of plant collections in Victorian times, and the changing methods usecl by thc star in caring for the important collections of plants. The rcinuiiiing scvcn chaptcrs prcscrit tlic struclure of the rainloresi and explain how the Palm lloiise exhibits it, llctailed discussions of the rainCOrest plants grown are presented in chapters on palms, cycads and pandans, tropical climbers, fruits, flowers, and plants of the forest floor. The section on palms is 23 pages long, beautifully illustrated in color and black and white, and is a well done and very complete overview of the family. The treatments of the other plant groups are equally good. (Continued on p. 52)