r99rl HAHN: ACANTHOCOCOS t67

Principes, 35(3), IS91, pp. 167-l7l

Notes on the Acanthococos

Wnrmlt JeMns HIHN Dept. of , Uniuersity of Wisconsin, Madison, WI 53706

Several recent authors (Moore 1973; In the later publication(1903), Barbosa "trlbe'o Dransfieldand Uhl 1986; Uhl and Drans- Rodrigues subdivided the Coco- field 1987) havelisted the SouthAmerican ineae Bentham & Hooker (currently rec- cerrado palm genus Acanthococos Bar- ognized as Cocoeae Martius) into seven bosa Rodrigues as synonymous with the subtribes,Attaleaineae BarbosaRodrigues, more widespread.Acrocomio Martius with- Eucocoineae Drude, Barbosineae Bar- out formal transfer of the type species.In bosa Rodrigues, Bactrisineae Barbosa addition, little or no discussionof the rea- Rodrigues,Astrocaryineae BarbosaRod- sonsbehind this reduction nor of the rela- rigues,Manicariineae BarbosaRodrigues, tionships of the genus and its component and.Leopoldiniineae BarbosaRodrigues. specieshas been offered. Current work on Manicariineae and.Leo poldiniineae are the of Paraguayanpalms (Hahn presently placed as subtribes in the tribe 1990, in prep.) has required an exami- Areceae,Attaleaineae (: Attaleinae nation of the genusand its three described Drude) is currently recognizedmore or less species. intact as a subtribe in rhe Cocoeae, and The type species,Acanthococos has- Barbosineaehas been united with Euco- sleri, was described by Joao Barbosa coineae as subtribe Butiineae Saakov Rodriguesin 1900 basedon a collection (Dransfieldand Uhl l986). of Emil Hassler from Ype-hu in eastern The two remaining subtribes, Bactri- and illustrated with a colored sineae(with Acanthococos,, Gui- plate in Barbosa Rodrigues' Sertum Pal- lielma Martius, Martinezia auctores (: rlarum Brasiliensium (1903). In the orig- Willdenow),Amylocar pus Bar- inal publicationoBarbosa Rodrigues spec- bosaRodrigues, and DesmoncusMartius) ulated about the affinities of the newly and,Astrocaryineae (with described taxon concluding that it was Martius and. Astrocaryurn, G. F. W. intermediatelo Bactris N. J. Jacquinand Meyer), are distinguishedfrom the others Syagrus Martius (under the then broadly by the presence of spines on some or all defined Cocos L.) citing the presence of parts of the . These are currently spines,free petals in the pistillate flowers, united as subtribe Bactrid.ineae J. D. and what he took to be basal endocarp Hooker (Dransfieldand Uhl l986). pores. This latter feature is anomolousfor No clear characters to differentiate the a spiny member of the Cocoeae.The non- two spiny subtribesfrom each other were spiny members of the tribe are variable in presentedin Barbosa Rodrigues' subtribal pore placement with a basal position most descriptions nor in his analytical key frequent (Uhl and Dransfield 1987). An (1903). Extracting informationfrom Bar- examination of material from the type bosa Rodrigues' generic descriptions, it locality and numerous additional speci- would appearthat the Astrocaryineae are mens from throughout the range of Acan- separatedby the staminate flowers sunken thococosohowever, shows an equatorial in pits on the rachillae accentuated by a rather than basalposition for the endocarp rim of raisedfloral bracts.This character, "honeycomb- Pores' which has beendescribed as I68 PRINCIPES [Vor. 35 like" in appearance (Uhl and Dransfield (1908), and Lem6e(1929, l94l) who 1987), is clearly illustrated in the color included the genus in the otherwise non- plate of Acanthococos hassleri (Barbosa spiny subtribeAttaleinae. Rodrigues, I903) though no mention of it Toledo differentiated Acanthococos is made in the various descriptions of that from genera of subtribe Bactridinae with taxon. The Cuban endemic Gastrococos sympetalousstaminate flowers (Bactris, Morales was not mentioned by Barbosa Guilielma (: Bactris), and' Astryocar- Rodrigues though it also bears the same yum) stating that it occupies a position characteristic rachilla pits. intermediateto theseand the choripetalous A second species from the cerrado of genera of the subtribe (Acrocom'ia, Braz'I, Acanthococos sericea, was de- Aiphanes, andDesmoncus). He calledinto scribed by Burret (1940) based on differ- question the subtribal separation of the ences in tomentum and spine density, spine spiny Cocoeae into groups by Barbosa color, and subterranean stem thickness. Rodrigues pointing out the similarities The author noted the similarities of Acan' betweenAcanthococcos and Acrocomia thococos with Bactris but differentiated it and between and Bactris. from that genus based on the free petals No mentionwas made of the choripetalous of the pistillate flowers. No mention was Gastrococos. made of endocarp pore position nor of the Wessels-Boer(1965) mentioned the rachilla pits. possibilitythat Aconthococosis synony- Toledo (1952) described a third species mous with Acrocomia but did not make from Brazil, Acanthococos e/nensis, with the new combinationdue to a lack of access two varieties, one each from the States of to material. Sao Paulo (var. emensis) and Minas Gerais Moore (1973) merely listed the name (var. pubifera), based on differences in as synonymous with Acrocornia without stem thickness, spine density, length of additional justification. In Genera Pal- staminate flower pedicels, and apparent tnarunl(Uhl and Dransfield,I9B7), a work differences in anther shape. His under- basedon Moore'snoteso Acanthococos was standing of previously described species informally placed in synonymy under was based entirely on published descrip- Acrocomia. Justifi.cationfor this was that tions and illustrations. An examination of the only difference between the two was specimens shows that stem thickness and in habit (p. 5I9). The genericdescription spine density tend to be variable characters of Acrocomia in Genera Palrnarum in both Acanthococos and the widespread includeslcan thococosand states an equa- (Jacquin) Lodd. ex torial position for the endocarp pores. In Martius. In addition, basal staminate flow- the subtribal description and in the key to ers are pedicellate in both of these taxa genera,however, a basalpore positionfor while the apical staminate flowers are pre- Acanthococos(under Acrocomia) is main- dominantly sessile. tained. Toledo (1952) also discussed the taxo- Generic boundariesin the Bactridinae nomic position of the genus correctly are fairly well established(Uhl and Drans- describing the illustrating of the equatorial field i987, p. 516) though the relation- rather than basal position of the endocarp ships of the various genera to each other pores. Based on this observationohe placed is still subjectto debate.A basicseparation the genus in subtribe Bactridinae th:us of genera into choripetalous and sympet- correcting the erroneous arguments offered alous generaas per Toledo (1952) would by Barbosa Rodrigues and perpetuated by seemto have considerablevalue though it Dalla Torre and Harms (1907), Pilger is difficult to apply to intermediate genera r99ll HAHN: ACANTHOCOCOS r69

and abun- such as Gastrococosand Aiphanes. The staminodialtube to the corolla,.With finely dividedand inconsistentsystem pro- dant, short mesocarpfibers. the cor- posedby BarbosaRodrigues (1903) relies rection of the erroneous original obser- too heavily on similaritiesin grossstructure vation of endocarp pore position, the large and ignores many of the detailed features number of similaritiesbetween the two gen- held in common. era justify placementof Acanthococosin InGenera Palmarum, no subdivisionof synonymy under Acrocomia. the spiny Bactridinae is employedthough Due to the abundance of edible and the genera are presented in a sequence industrial quality oil in the mesocarp and of proposed evolutionary advancement. endosperm(Balick 1979, Lleras 1985), Reduction in number of pistillate flowers the genusAcrocomia hasbeen the subject and connation of perianth parts, especially of considerablebiosystematic and evolu- in the pistillate flowers, are characteristics tionary study (e.g., Scariot l9B7). The considered most important in assessing taxonomy of the genus, howeveroremains specialization. uncertain(Uhl and DransfieldI9B7). Ear- The genus which has been most fre- lier workers (i.e., Bailey 1941, Glassman quently associatedwith Acrocomia, Gas- 1982) maintained the genus with a large trococos, shares a similar floral arrange- number of speciesmostly basedon natrrow, ment, rachilla chambers, and a connate typological definitions. Recent work con- staminodial ring adnate to the corolla but cerning the systematics of neotropical can be differentiatedby the connate sepals palms, however, supports rather broad of both staminateand pistillate flowers,the speciesconcepts for many groups,partic- pistillate petals distinctly connate basally, ularly those which have been exploited by the erect filaments with basifixed anthers, man or which occur in environmentsinflu- and the absenceof short mesocarp fibers enced by local human populations (Balick adherent to the endocarp. The only other et al. l9\7a" b: Anderson and Balick choripetalous genus in the subtribe, l9BB). As the genusAcanthococos occurs Aiphanes, shares rachilla chambers, free in areas of {requent fire or tilling (Eiten sepals, and a staminodial tube adnate to 1963, I972), these observationssuggest the corolla, but differs in its praemorse that the three describedspecies of Acan- leaflet margins, erect filaments with basi- thococos are merely minor deviations of fixed anthers,basally connate pistillate pet- but one, somewhatvariable species. als, and absenceofmesocarp fibers. In the The northeastern corner of Paraguay sympetalousAstrocarlrum, similar rachilla has produced a number of unusual and chambersand mesocarpfibers are present apparently endemic taxa and is without but the pistillate petalsare connatebeyond doubt the most floristically diverse region the middle and the staminodial tube is very ofthe country. Severalofthese taxa, how- small and not adnate to the corolla. ever, were based on incomplete material Characteristics shared by both Acan- or inaccurate interpretation of structures thococos and,Acrocomia include a floral as in the caseof SchrankiastnrnzHassler arrangement with a few, solitary pistillate (Barneby1984). A similarsituation appears flowers at the base with numerous paired to be present it Acanthococosin which or solitary staminate flowers toward the the misinterpretation of endocarp pore apex, distinctive rachilla chambers, free position led to an improper taxonomic sepalsof both staminateand pistillate flow- alignment of the genus. After clarification ers, filaments inflexed in bud with dorsi- of this problem, lack of additional work on fixed anthers, free or only very partially thisand relatedgenera left properaffinities connate pistillate petals, adnation of the obscured.Overemphasis of relatively minor I70 PRINCIPES [VoL.35 variations in morphology and maintenance C. U. B. PINHEIR0e.no A. B. ANoeRSon. I987a. Hybridization in the Babassupalm com' of typological species concepts led to plex: I. Orbignya phalerata x O. eichleri. inflated taxonomies and further obscured Amer. J. Bot.74(7): I013-1032. taxonomic relationships. A. B. ANDERSoN.eNo J. T. un M.torr- In light of the above arguments, I make nos-Coste. 1987b. Hybridization in the x the following new combination and unite Babassu palm complex: Attalea compta Orbignya oleifera (Palmae).Brittonia 39(1): 26- synonymy the two species of Acantho- in 36. cocos described after the type species, l. BaReose RoDRIcUES,J. 1900. Palmae Hassler- hassleri. ianae Novae uo Relacao das Palmeiras encon- tradas no Paraguay pelo Dr. Emilio Hassler de I898-1899. pp. t I3. . (BarbosaRodrigues) 1903. Sertum Palmarum Brasiliensium, Hahn,comb. nov. ou Relation des Palmiers Noveaux de Bresil, Acanthococos hassleri Barbosa Decouverts, Decrits et Dessinesd'apres Nature. Rodrigues, Palmae Hasslerianae 1: l-140.91 plates;2:I-114,83 plates.Veuve Monnom. Bruxelles. Novae2. 1900. Syntypes:Paraguay, BARNEBY,R. 1984. A new Bolivian Mimosa of Ape-hu Hassler 4957, 5224 (pre- seclion Habbasia series Leptostachyae (Legt- sumably destroyed).Neotype: Para- minosae:Mimosoideae), close kin of the fictitous guay, Yhu (Holotype:G). genrs Schrankiaslrum Hassler. Brittonia 36: 248-251. Acanthococos sericea Burret, Notiz- BunRnr, M. 1940. Eine neueArt der bishermono. blatt 15: 109. 1940. Type: Brasil, typischen Gattrrg Acanthococos B. R. in Mat- Mato Crosso.Archer & Gehrt36432. togrosso gefunden. Notizblatt Bot. Card. Berlin (Holotype:SP; isotype:US). 15:109. 1907. Acanthococosemensis Toledo, Arqui- Derr.t Tonnr. C. G.. Dn AND H. HARMS. Genera Siphonogamarum,Suppl. p. 592. vos de Botanica do Estado de Sao DRANSFIELD,J. eno N. W. UHL. 1986. An outline Paulo,Nova Serie 3: 4,t. I-2.1952. of a classificationof palms. Principes 30(l) 3 Type: Brasil, Sao Paulo. M. Rachid 1I. 52, 941. (Holotype:SP). EIrtN, G. 1963. Habitat of FazendoCam- pininha, Simposio sobre o cerrado. p. 216 Edi Acanthococosem,ensis var. pubifolia toria de Universidade de Sao Paulo. Toledo,Lc. 3: 5. I952. Type: Brasil, 1972. The cerrado vegetation of Brazil. MinasGerais. L. Labouriau 53.727. Bot.Rev. 38: 201-341. (Holotype:SP). Gr-,tsslreN,S. F. 1982. A revisionof B. E. Dahl- gren's Index to American Palms. Phaneroga- The speciesis distributed sporadically marum monographiae.J. Cramer. Lehre. throughout the Planalto of Central Brazil He.Hu,W. J. 1990. A Revisionof the Palmaeof M.Sc. Thesis,Cornell University, Ith- from the Sierra de Amambay in north- Paraguay. aca, New York. easternParaguay to westernBahia and is in prep. Palmae. In: R. Spichiger and J. locally common in cerrado vegetation. M. Mascherpa(eds.), Flora del Paraguay'Geneve' (Medeiros-Costaand Panizza I 983). LEM6E,A. I929. Dictionairedescriptif et synony- mique des genres de plantes phanerogames, I: I5. f94I. ibid. 8a: 86. (in analyticalkey) LlrnRerunn CIrnl LrEnqs, E. 1985. Acrocomia.um genero com grandepotencial. Noticiario 1: 3-5. EMBRAPA,U ANDERSoN,A. B. eNn M. J. BALICK. l9BB. Tax- CENARGEN, Brasilia. onomy of the Babassucomplex (Orbignya spp.: MEDETRos-Cosre,J. T. or eNo S. P,tNIzze. 1983. Palmae).Systematic Botany I3(I): 32-50. Palms of the Cerrado formation of Sao Paulo BAILEY, L. H. f94I. Acroconia Preliminary State, Brazil. Principes27(3): I18-I25. Paper. GentesHerb. 4(I2): 421-476. Moonr, H. E., Jn. 1973. The major groups of BALICK,M. l. 1979. Amazonianoil palmsof prom- palmsand their distribution.Gent. Herb. 11(2): ise: a survey. Econ. Bot. 33(l): 1l 28. 27-r4t. r99rl NEWS OF THE SOCIETY

PILcER, R. 1908. Palmae. In; A. Engler & K. thococosBarb. Rodr. Arq, Bot. EstadoSao Paulo Prantl, (eds.),Die Naturlichen Pflanzenfamilien, 3: 3-5, t. I-2. Nachtr. III z. 2-4 Teil, p. 27. Uur, N. W. erqo J. DReNsntnrl. 1987. Genera ScARIor, A. O. 1987. Biologia reproductiva de Palmarum. A classification of palms based on Acrocomia aculeata (Jacquin) Loddiges ex Mar- the work of Harold E. Moore, Jr. L. H. Bailey tius no Distrito Federal. Master of ecologythesis. Hortorium and The International Palm Society. Universidade de Brasilia. 610 pp. ToLEDo, J. F. 1952. Estudos sobre algumas Pal- WESSELSBonn, J. G. 1965. The Indigenous Palms meiras do Brasil. II. Notas sobre o generoAcan- of Suriname,E. J. Brill, Leiden,pp. I 172.

Principes,35(3), I99I, pp. l?1-179

CHAPTERNEWS AND EVENTS

New Journalfor TemperateZone NEWSOF THE SOCIETY Enthusiasts:The Hardy Palm The IPS has lost three well known and The Pacific Northwest Palm and Exotic influential members during the past few Plant Society, an affiliated chapter of the months. Obituaries for one of them, Hans IPS, has resumedpublication of a Chapter Hoffmann, is given here. We will have Newsletterto meetthe needsof Temperate articles on Ruth Shatz and Dr. Anthony Zone palm enthusiasts.The Hardy Palm Davis in October.-eds. carries articles of interest to palm hob- byists in colder regions. The editor is Nick Parker. The Hardy Palm averages16- PacificNorthwest Chapter Meetings 24 pages(5.5 by 8.5 inch size)and will be issuedquarterly. Interestedparties from The Pacific Northwest Chapter of the other cold weather elimatic zones are IPS held their SecondCeneral Meeting for invited to subscribe.Subscription rates are I99I on Tuesday,May 28, l99l at the US $ 12 per year for United States IPS VanDusen Gardens in Vancouver, B.C., membersand Cdn $ 15 per year for Cana- Canada,with guest speakerGert Vander dianmembers and Cdn $ I 9 (or US $ I 5.00) Meulen discussingthe latest in greenhouse for mailingsoutside North America. Checks construction.The Third and Fourth l99l in US or Canadiancurrency payable to GeneralMeetings will be held on Mondays, The Pacific NorthwestChapter, IPS, should August 26 and November 25, respec- be sentto Frank Hanaus(Canada) or Nick tively, at the same venue. The group's Parker (Canada).Articles and materials third annual Palm Society BBQ will be held suitable for publication are also solicited in July. Also plan to visit the Palm Society and should be sent to Nick Parker, Editor Booth at the PNE betweenAueust l8 and {seeroster). September2, 1991.