19771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 47 -The Needle Palm: Rhapidophyllum

Ar-r-riv G. Suunv AND RIcHARDP. Wulvnrnr,ru Conseraation Consuhants, Inc., Palmetto, 3356I; anil Depa.rtment of Biology, Uniuersity ol South Fhrt(a, Tampa, Floida 33620

Rhapidophyllwn is a monotypic palm seed is planted until a marketable genus endemic to the southeastern is produced, thus not making it eco- . Its single species, R. nomically febsible for nurseries to grow hystrix (Fig. I), is commonly referred it from seed. to as the needle palm. Other names Rhapid,ophyllunz is on the rare and which have been used are blue pal- endangered plant species list for the metto (Elliott, 1817), creeping palmetto United States as a threatened species (Small, 1923), vegetable porcupine which is commercially exploited. Dur- (Small, L923), dwarf saw palmetto ing the late I800's and early 1900's, this (Moore, 1963), and spine palm (Mitch- palm was commercially exploited by ell, 1963). foliage shippers. Whole crowns were In the United States, the needle palm cut off and shipped north for use as is cultivated to a limited extent both decoration in homes. This has been within and outside its natural range. called to our attention by Harper It is very cold hardy, having been suc- (1906) who lamented the exploitation cessfully grown out of doors as far north o{ this pabn in the vicinity of Ever- as Virginia arl0 (Popenoe, green, . Later, Harper (1928) L973). related that the species has been ex- Outside the United States, R. hystrix terminated in the Evergreen area. To- has been successfully grown in the day the exploitation continues, but is Black Sea region of the Soviet Union through the nursery trade. (Saakov, 1963) and on the French Riv- This paper is for a large part the iera (Barry, 196I). result of the senior author's personal Although Rhapid,ophyllum has gteat observations of both wild and culti- potential as an ornamental because of vated specimensover a five-year period. its attractiveness, ability to withstand Field observations were made through- Ireezing temperatures, and ease of out much of the range of the sptjcies maintenance, it is rare in the nursery in Florida with the more critical and trade. Only recently has it begun to detailed observations made on a pop- to be used in public and private land- ulation near Oviedo, Seminole County, scaping. Most found in the nurs- Florida. ery trade and in cultivation have been taken from the wild. This is probably Tqxonomic History due, at least in part, to the fact that Rhapid,ophyllum,is a monotypic genus seeds are slow to germinate and the of coryphoid palms in the Trithrinax plant is very slow growing. It takes unii of the Trithrinax alliance (Moore, from {our to six years from the time L973). This unit also includes the gen- PRINCIPES lVor.. 2l

l.Rhapid,ophyllum hystrix near Oviedo, Seminole Counry, Florida.

s era Tra,chycarpus and, Chamaerops in ingly created the monotypic genus ,Rfra- addition to Trithrinax and Rhapid,o- pidophyllum. phyllwm. Small (1923) states that the species Small (1923) presenteda fairly com- has been placed in two other genera in plete and accurate taxonomic history of addition to those previously discussed, Rhapid,ophyllurl so only a few points Corypha and. Rhapis, although he does need to be touched on here in summa- not elaborate. tion and for clarification. Dahlgren ( 1936), and later Glass- In 1814, Frederick Pursh described man (1972), give Rhapis arund,inacea the speciesfrom material collected near Ait. (Hort. Kew. ed. I, 3: 474. l7B9) Savannah, Georgia by John Fraser sev- and.Rhapis caroliniana Hort. ex Kunth eral years earlier, and placed it in the (Enun. Pl.3:246. l84l) as synonyms genus Chamaerops. In lBlB, Thomas oI Rhapi.d,ophyllum hystix. From Ai- Nuttall, apparently having seen only ton's protologue of Rhapis qwndina,cea, sterile material, transferred it to the it is immediately obvious that the plant genus Sabal because of its superficial described is not Rhapid,ophyllum hys- resemblance to S. ad,arconii (= S. mi- trix. zor). Finallyo in L876, Wendland and William Wood (ln Rees, Cycl. 7, n. Drude found that the species could 3. fB07) transferred Aiton's speciesto not be satisfactorily accommodated in Chamacrops with no additional clues either Chamaerops or Sabal and accord- to its identity. I9771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 49

Finally, Moore (W75), after having to Pursh by some authors is an error examined the type spbcimen oI Rhapis in citation. arund,inacea in the British Museum In summation, the following nomen- (Natural History), reveded Aiton's spe- clature is to be applied to the needle cies to be based on an a:berrant,juvenile palm: specimen of. Sabal rninor. Rhapidophyllum hyetrix (Pursh) H. Also, Rhapis cololiniaw, the second Wendl. & Drude, Bot. Zeit. 34: 803. Rhapis species placed in synonymy un- L876. der Rhapid,ophyllum hystrix by Dahl- Chamadrops hystrix Pursh, Fl. Amer. gren and Glassman, can also be ex- Sept. I: 240. I8I4. cluded. The name Rhapis cq,roliniq,na Sabal hystrix (Pursh) Nutt., Gen. N. is not found under the discussion of Amer. Pl. 1: 230. 1818. Rhapis which appears on pages 251- 252 oI Kunth (1841). However, on page 246 under the treatment SaboJ Hqbitqt qnd Disiribution (= adarwonii S. minor), the name Sa- Rhapid,ophyllum, hystrix is an un- proposed bal carolinianurn Hort. is in common speciesnative to the southeast- synonymy. If this is the source of ern United States, occurring from Beau- Rhapis caroliniana Hort. ex Kunth, fort County, , south to which appears to be the case, this name Hiehlands and Hardee counties, Flor- the must be removed from synonymy ida, and west to Simpson County, south oI Rhapid,ophyllum hystrix and added central (Fig. 2). Distribu- Sabal rninor. to that oI tional data are compiled from herbar- validly published name in No Cory- ium specimens, literature, and sight pha can be accredited to R. hystrix. records (pers. comm.). Questionable hystrix DesI. (Tabl. l. Corypha 19. records in the literature which could 1804) is a nornen nud,um and in addi- not be verified with specimens are ex- tion undoubtedf not synonymous with cluded. Rhapid,ophyllwrn hystrix since it is re- Throughout its range, Rhapid'ophyl- ported as a native of South America. Ium is found primarily in low, moist to hystritc (spelled "fris- lilso, Cha,tnaerops wet sites with rich, humus, calcareous (Nom. tix" by Steudel) Desf. ex Steud. clayo or sandy soils in woods, swamps, Bot. ed. 1. I83. 1840. based on Corypha and hammocks (Fig. 3). Less com- hystrix Desf.) is also a nornen nud'um monly, it occurs in limestone sinks, and is accordingly removed from con- grottos, and shaded pinelands. Al- sideration here. though rarely found in full sun and Martius (f$B) provides an excellent well-drained sites in the wild, the spe- description and illustration of the needle cies will thrive in such locations in cul- palm under Chamaerops hystrix. In tivation if sufficient soil moisture is present. addition to Sabal hystrixo Rhapis arun- Specimens occasionally found in the wild on well-drained sites are d,inacea, and Charnaerops arund,inncea, generally smaller and in poorer con- Martius also lists Corypha reperls Battr. dition as compared to those found in in synonymy. The last is now well wetter soils. There does not appear to (Bartr.) known as Serenoa reperls be a specific correlation of soil type Small. witlr the presence oI Rhapid,ophyllurn. Of final note, the crediting of. Cha- However, in the north central part of nxa,eropshystrix to Fraser instead of its range it is generally associatedwith 50 PRINCIPES [Vor. 2l

a aaa aa O a o oo o

2. Distribution of. Rhapidoplryllum hystrix. limestone soils. In central Florida. Hurricane Camille in 1969 was a large where the limestone bedrock is over- and beautiful clump in 1973. It ap- lain with sandy soils, Rhapid'ophyllum parently was not severely affected. often occurs in seepageareas along the Rhapidophyllum can be considered edges of the various uplands constitut- locally common or even abundant in ing the central Florida ridge area. some areas within its range, but its Rhapid,ophylhum apparently has a distribution is irregular. It was ob- fairly high degree of salt tolerance, at served to be the dominant understory least for periods of short duration. To species in several Florida hardwood cite a specific example: a cultivated swamps, although it is rarely found specimen in the medial strip of US 90 where it would be highlv competitive near Biloxi, Mississippi, which undoubt- with other species. edly was submerged in salt water by The species does not appear to be r9771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 51

3. Rhapid,ophyllum colony near Oviedo, Seminole County, Florida.

I occupying all- the available suitable annually. The reason for this irregu- habitat. Many areas that appear en- larity is not known, but may be due vironmentally identical from visual ob- in part to less than optimum environ- servations to those occupied by Rha- mental conditions. For example,- pidophyllwn do not support the species, ing in central Florida populations in even though the occupied and unoc- 1970, a wet year, was good, but since cupied areas are often found quite near 1970 the seasonshave beenconsiderably one another. Also, what appears to be drier and as a result flowering less fre' a uniform habitat is frequently poorly quent and more irregular. However, utilized by the species. Some possible cultivated specimens which appear to reasons for this irregular distribution receive sufficient moisture will also will be discussedlater in this paper. flower irregularly. Although the avail- ability of moisture may play a role in Reproduciive Biology initiating the process, flowering ap- Flowering generally occurs from pears to be physiologically more com- March to August with flowering having plex. been observed in both cultivated and Rhapidophyllzrrz is usually dioecious, polygamodioecious wild plants as early as February and less commonly or as late as November. Flowering in. the rarely monoecious. In most popula- wild is irregular and infrequent' In- tions staminate or predominately sta- dividual plants generally do not bloom minate inflorescences are more fre- 52 PRINCIPES lVor-.21

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4. Specimen showing staminate inflorescence. 19771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 53

6. An unusual -carpellate flower (left) and a typical 3-carpellate flower (right).

times reduced to one or two by abor' 5. Close-up of pistillate . tion or rarely increased to four (Fig. 6). This latter condition is highly un' usual in palms and merits further in' quently observed than pistillate. The vestigation. The young carpels are yel' staminate inflorescences (Fig. 4) are low to yellowish-green, turning yellow the more conspicuous of the two with or yellowish-cream at anthesis. The most or all of the inflorescence being rachis of the pistillate in{lorescence is forced clear olrthe bracts. The pistillate purple. After pollination, or in age, inflorescences are less noticeable, being the corolla turns a whitish or yellowish much more constricted and not well color, eventually becoming brown. The projected from the bracts with half or four to six bracts are cream or dirty more usually remaining permanently white in color, occasionally purple- enclosed. A close-up view of the pis- tinged. Two major color variations of tillate flowers is seen in Figure 5. pistillate flowers were observed in the Flower color has been reported as be- Seminole County, Florida populations. ing yellow, orange, or purple. We have Two plants were seen which had a clear observed all three colors in the field citrus-orange corolla and a whitish with purple predominant. Staminate calyx. The rachis of the inflorescence {lowers appear yellowish from a dis- was purple. A third plant was observed tance because o{ the predominantly with an orangish-purple corolla and a cream-colored anthers. The calyx, co' purplish-white calyx. The rachis of this rolla, and rachis vary from a dirtY in{lorescence was also purple. These cream-color to pale pink to deep purple {lowers were fresh, having emerged from or brownish-purple.The corolla of the the bracts only a short time before ob- pistillate flowers ranges from brownish- servation. Although these color varia- purple to deep purple before pollination. tions are striking, it is improbable that The gynoecium in Rhapidophyllum is they have any taxonomic significance. apocarpus and usually tricarpellate. During the day, the staminate {lowers However, the carpel number is some- emit a distinct musky odor. The pistil- 54 PRINCIPES lVol-. 2l late flowers give o{f little or no odor are usually produced on well-projected during the day. However,'produced they emit a inflorescenceswhich resemble staminate weak version of that by the inflorescences.The reason for this con- staminate flowers during the evening dition is unknown. hours. It has not been determined if Rhapid,ophyllwn seeds are extremely a fragrance is produced by either the well protected by spines surrounding pistillate or staminate flowers at night. the trunk. They appear to be largely The exact time relationship of the pol- unattractive to animals since they are linator visitation needs further inves- rarely disturbed. The pistillate inflores- tigation. cence is so short that most are Rhapid,ophyllum is self-compatible caught in the cluster of petioles and since isolated, cultivated plants will oc- needlessurrounding and below it. Most casionally set abundant . As the o{ the fruits remain in the crown until fruit matures, the white-hirsute condi- they either decay or sprout. Manley tion, which is very evident when young, (1967) found the ripe fruits of needle becomesmore diffuse, but is persistent. palm chewed on by what he thought The surface of the mature fruit varies to be mice. Rhapid,ophylhrnz seeds in from brown to reddish- or purplish- the Seminole County, Florida popula- brown to red. The flesh is orangish and tions have been found scatteredaround somewhat mealy in consistency. The the bases of the parent plant. Most of fruits mature from December to Feb- the flesh had been removed and the ruary. The pistillate inflorescences re- seed bore marks produced by the teeth main crowded in the bracts and sheath of a small mammal. The fresh mature fibers during fruit maturation. As a fruit apparently has few properties to result, the fruits on the lower and inner attract animals. It is bland in taste. parts of the stalks usually become being only slightly sweetish. Little or greatly distorled (squarish or pyriform) no odor was detected. However, over- while those 6h the upper part possess ripe fruit emits a strong, overly sweet, the more characteristicovoid or globose pungent odor. Manlev Q967) describes shape. The fruits average 1.5-2.0 cm it as being like rotten cheese. It may in diameter. Fruit clusters weighing up be the odor of the overripe fruits that to 845 grams have been found. Most attracts animals. This odor is appar- fruits possess a small spine near the ently produced after the fruits have been apex of the endocarp. This spine varies subjected to freezing temperatures. from practically nonexistent to 2 mm The immature" half-formed fruits in length. The endocarp frequently has seem to be preferred by some animals a marking resembling that of pressed over the mature fruits. Several times cork board. However, some endocarps, the remains of immature seeds were are smooth. observed scattered around and near a Inflorescences well projecting from fruiting palm. It appears that what- the subtending bracts and resembling ever feeds on the fruit, possibly squir- the staminate, but with scattered fruits, rels, mice, or wood rats, prefers the en- are sometimes encountered. This is in- dosperm before it becomes hard. It is dicative of the polygamodioecious con- also noted that the flesh of the green dition. Plants have also been observed fruit is very astringent in taste, similar that produce abundant fruit which is to that o{ a green persimmon. smaller than normal and hollow or with In addition, many fruits are parasit- shriveled" inviable seeds. These fruits ized by small cereal beetles which make r9771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 55

small burrows into the hard endosperm plants tend to grow away from each making an opening through the endo- other. In time the connection between carp of the fruits which have the exo- the parent plant and the offshoot is sev- carp and mesocarp removed. These ered by the rotting away of the stem openings permit the easy entry of fungi tissue, resulting in the establishment of and other decomposing organisms which two individual plants. further damage the seed so that it even- Many Rhapidophyllum specimenshave tually becomes inviable. As a conserva- a slanted or decumbent trunk with three- tive estimate, about 20% arc parasitized fourths oF more of it covered with a by these beetles. thick coat of spines and fiber. The re- Seed germination takes from about mainder o{ the trunk is naked with rings six months to about two years. The of previous bases quite evident. iirst divided appear about three Palms with upright trunks or those in years after germination. Small, weak dense clumps generally have spines and spines are evident at this time. Seed- fiber almost to the ground. In large lings were found to be uncommon in plants a sizable cloak of leaf bases and the wild on the ground; however, many fiber is retained on the trunk. The were found in the crowns of the parent trunk may appear to be 0.5 m or more plants where they germinated among in diameter although it actually may be the spines and thick dark brown fiber only &-10 cm. The spines are the result surrounding the trunk. The chances of the partial breakdown of the leaf of these seedlings surviving is low be- sheath tissue so that only the rigid cause of the lack of available nutrients sclerotic tissue persists (Tomlinson, and water. The fiber and spines pro- L962), The spinesrange in length from duce a favorable environment for few 24 dm. The retention of the leaf base plants other than some fern species. fiber prevents the loss of the spines. It Only one Rhanid,ophylhrm population may be speculated that the spines prob- was found in Florida (Stallion Ham- ably evolved to protect the rather suc- mock, Hillsborough County) where a culent flowers from herbivores rather pronounced increase in population by than the growing tip of the palm. Seed- seedlings could be detected. The seed- lings four or five years old produce lings were well establishedon the ground only weak spines and if spines were and outnumbered the adult plants. The needed for the protection of the grow- other populations examined usually did ing tip, they would develop early in the not have enough seedlings to replace life history of the plant. There may even a small percentage of the existing have been strong predation pressure population o{ palms. causing the palm to develop the spines One offset to poor production of vi- and constricted in{lorescences. A sec- able seed in Rhapid,ophyllum is that it ond reason for the leaf base and fiber is relatively long-lived, allowing it to retention is for the protection of the produce seedover a long period of time. roots. It seemsthat the reclining trunk Another factor in the maintenance of of most Rhapid,ophyllum in the wild populations is that the species repro- is not the result of a natural tendency to duces by suckering. Although this does lean or crawl, as it is in some speciesof not increase its range at a rapid rate, it palms, such as Sabal etonia, but be- does increase its numbers within the cause of unstable anchoring. Seedling population. Rhapid,ophyllum show no tendency to Offshoots produced by suckering lean as do seedlings of Serenoa and, JO PRINCIPES lVor.21

Sabal. As the needle palm grows, the cences or suckers. Of the buds pro- end of the trunk iots off, taking with duced, about 50% abort and o{ the re- it any roots which would give support. mainder, B0% produce inflorescences In time, the weight' of the palm or and only abot 20% produce suckers. other forces causesthe palm to tip or Also suckering may be inhibited by a fall over. Rhapid,ophyllum has the abil- strong, healthy apex. Until there is dam- ity to produce roots anywhere along the age to the apex or su{ficient distance trunk, no matter what age the plant or created by addition of the trunk be- the diameter of the trunk. A large palm tween the apex and the vegetative bud, examined from the Seminole County suckering will be inhibited. This inhi- populations showed numerous roots bition may also be effective in Rhabido- about three-fourths the way up on all phyllum and would account for the ab- sides of the trunk B cm in diameter. senceor sparsesuckering in old, large, The roots were growing among the old single-stemmed plants with a large leaf bases and fiber, damp and well crown. However, there may also be protected. The cloak of fibers there' other explanations such as environmen- fore acts not only as an initial protec- tal restrictions. A single-stemmedplant tion for the young roots, but as a damp taken from the wild will often sucker medium for the roots to pass through profusely, probably due to transplant from the trunk to the soil should the shock. It is also of interest that seed- palm fall over. It also is conceivable lings, to the best of the authors' knowl- that the young roots function as absorp- edge, always produce suckers. tion organs, imbibing water from the The ability of the plants in cultiva- spongy trunk mantle. On one occasion tion to grow taller is probably due to a specimen was observed with about a better growing conditions and more foot of trunk covered to ground with light. Drier conditions and probably fiber and spi-nes.Upon examination, it better air circulation keep the trunk was found tfiat only one root reached and roots from rotting. The creeping into the soil. The rest of the functional habit which Rhapid,ophyllum in the wild roots were found intermeshed in leaf exhibits has rarely been seen in culti- bases and fiber. vated plantings. Roots from the canopy and under- Rhapid,ophyllum is not vigorous or story trees often invade the fibrous aggressive and it is doubtful whether it mantle ol Rhapid'ophyllum. These roots could successfully compete with helio- may eventually provide support for the philic plants. However, its slow growth poorly anchored palms. probably allows it to occupy habitats Rhapiilophyllum ilso has the potential with low light intensity. As suggested to produce suckers anywhere along the in a study oI Iguanura geonomilormis trunk regardless of the size or age of (Kiew, L972), the success of under- the plant. Suckers may be found on growth palms in occupying a dark habi- di{ferent plants from the base of the tat is due to their slow growth because trunk to within several inches of the not much light is required to maintain growing apex. Still, many palms {ail them. Rhapid,ophyllunr, like several to {orm suckers. Fisher and Tomlinson other species o{ palms, apparently is (1973), in their studies of the branch- occupying a habitat where it does not ing suckering oI Serenoa repens, Iound. have to compete with faster-growing that this species produced an axillary plants for light, moisture, and nutrients, bud which could either produce inflores- Bannister (1970) noted, in the case o{ r9771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 57

Prestoea montana, (reported as Euterpe inflorescences of such diverse species globosa) in Puerto Rico, that it is most as Albizzia sp. (Fabaceae), Asimina likely that seedlingsof this, and perhaps paraiflora (Annonaceae), Cassia tora other forest speciesas w6ll, may be able (Fabaceae), Castanepmollissima (Fag- to exist in a state of semidormancy until aceae), Carica wpaya (Caricaceae), some environmental factor becomes Gossypium hirsutum (Malvaceae), 1/i- more favorable, at which time rapid biscws sp. (Malvaceae), Nerium ole- growth can resume. A similar'responso and,er (Apocynaceae), Passiflora sp. may also occur in Rhapid'ophyllum. In (Passifloraceae), and Pluchea loetida the case of Rhapidophyllum, as perhaps (Asteraceae). The larva has also been in other forest palm species as well, known to bore into the stem of sugar light is suspected to be the limiting cane (Saccharum officinarum-}oa- factor. Rhapid,ophyllum, when rrans- ceae). Notolomus has also been ob- planted from a dark to a well-lit site, served on the inflorescencesof Serenoa will usually grow much more rapidly. repens and. Forestiera segregata (Olea. Popenoe (1973) reported that the ceae). Based on the known preference specieshas withstood temperatures down of the two described species of this to 12 degreesbelow zero without being genus, it is probable that N. bicolor is damaged. The minimum temperatur€ the species found on Serenoa while ff. that Rhapid,ophyllum can withstand basalis is found on Forestiera, without being killed has not been de- This undescribed Notolorn&s species, termined. It certainly is one of the found in great abundance on the inflo- more hardy palms. rescences of. Rhopid'ophyllum, is the In April, L975, a survey of the insects prime candidate for pollinator. Adult on the flowers oI Rhapid'ophyllum was beetles taken from the staminate in- made. Flowering at that time was poor florescences were found to be heavily and a high proqortion of staminate in- covered with pollen. Notolomws sp. nov. {lorescenceswai-produced as compared is ideally suited as a pollinator since to pistillate. it is small, active, and a strong flyer. Inflorescences were collected with the When the inflorescence is disturbed, the staminate and pistillate placed in sep- beetles will respond by quickly drop- arate containers of alcohol for preserva- ping {rom the inflorescence into the tion. These were later examined and fiber or spines surrounding it, by crawl- the insect visitors noted. A weevil, ing deeper into the inflorescence, or by identified by Rose Ella Warner as prob- flying away. Since the compacted pis- ably a new speciesof Notolomus (Cur' tillate inflorescence does not emerge culionidae) near ff. basalis, comprised far from the bracts a small and active more than 9O% oI the insects found on pollinator would be necessary fior Rha- both types of inflorescences. Fresently pid,ophyllum. only two described speciesoI Notolom,us As previously stated, staminate flow- are known from North America: ff. bi- ers, and to a lesser extent, the pistillate color and. N. basalis. Based on data flowers, emit a musky odor that serves from the entomological collections at as the attractant for the beetles which the United States National Museum, 1V. then feed on the pollen and various bicolor has been collected on the inflo- flower parts. The large numbers of bee- rescencesoI Chamaerops sp. and Sabal tles present (estimatedto be about a hun- palmetto. Notolomus basalis, on the dred per in{lorescence),their small size, other hand. has been collected on the pollen load, and mobility make this in- JO PRINCIPES lVor 2l sect an effective.pollinator. It is pos- mon pantropical and pansubtropicalspe- tulated that the beetle is first attracted cies. These organisms probably account to the area by the strong odor produced for some of the damage to the flowers, by the staminate fl.owers, which usu- resulting in lower fruit set. -. ally outnumber the pistillate in a given The needlepalm appearsto be a senes- area. The beetle will then go {rom in- cent species. Flowering is irregular, florescenceto inflorescenceseeking {ood. {ruit set poor, seedparasitism high. Its Pollination by curculionid beetles is distribution is irregular and suggests a not rare in palms. Essig (I97I) pro- disintegrating range. Available habitat vides evidencethat in Costa Rica, two is also diminishing, due primarily to speciesoI Bactris are pollinated lry PhyI- human activity. There appears to be lotrox megalops and possibly Grasid,ium no effective long-range dispersal mech- longimanus along with two specieso{ the anism {or the establishment of new nitidulid beetle genus, Mystrops. Essig populations in now existing suitable (1973) also suggestedthat the curculi- habitats. Reproduction is apparepfly onid beetle Nod,ocnemus sp. pollinates primarily by vegetative means, which Hydriastele rnicrospad,ix in Papua New results only in the maintenance rather Guinea. This latter genus of beetles and than expansion of existing populations. related genera are believed to occur al- Expansion of populations apparently most exclusively in palm flowers and only takes place locally. Rhapid,ophyl- are pantropical in distribution (Essig, lum hystrix is best interpreted as a Le73). slowly vanishing relict species. Schmid (1970) observedcurculionids (Celestessp. and Phytotribus sp.) on Acknowledgments Asterogyne martiarw, in Costa Rica, but concluded that they were not effective Herbarium specimens were examined pollinators for a number of valid rea- from the following herbaria: ALU, sons. Similryly Brown (1976) noted AUA, CU, FLAS, FSU, GA, GEO, Notolomws basalis on Sabal palmetto, MISS, MISSA, NCU, NY, USF, ANd but concluded that this palm was pol- VSC. The curators of these herbaria linated by bees, primarily Apis mellif- are grate{ully acknowledged. Dr. Rose era, rarher than by this curculionid. Ella Warner, Smithsonian Institution, Various other organisms were found provided the determination for the in the inflorescencesoI Rhapid,ophyllum, curculionid beetle and is gratefully ac- but are discounted because they either knowledged. Special thanks are ex- were not found in suf{icient numbers or tended to Dr. Frederick Essig, Uni- were not physically able to satisfy the versity of South Florida, Tampa, for requirements for pollination of this spe- his critical reading of the manuscript cies, Among these were neuropteran and help{ul eomments. Others who pro- larvae, immature hemipterans, isopods, vided valuable information and are ac- arachnids, lepidopteran larvae, chilo- knowledged are: I7illiam D. Manley, pods, various coleopteran larvae, an am- Stockbridge, GA; Dr. Wayne R. Fair- phipod, and a snail. The amphipod, cloth, Valdosta State College, Valdosta, which was very abundant on both the GA; Dr. Edward L. Bousfield, National in{lorescences and in the leaf litter on Museumsof Canada,Ottawa; Dr. James the forest floor in one population of W. Hardin, North Carolina State Uni- Rhapi.d,ophyllum,has been identified by versity, Raleigh, NC; Dr. John Popenoe, Edward Bousfield as Talitroid,es topito- Fairchild Tropical Garden, Miami, FL; tum (Talittidae), an increasingly com- Dr. J. R. Watson, Mississippi State Uni- r9771 SHUEYAND WUNDERLIN:RHAPIDOPHYLLUM 59 versity, Mississippi State, MS; James 1928. Economic BotanY of Ala- part 2. Geological Survey oI Ala- E. Poppleton, University of South Flor' bama, bama, University of Alabama. Monog. 9. ida, Tampa, FL; Steven'W'. Leonard, Krew, R. 1972. The natural history ol lgua' Tallahassee.FL. The authors gratefully nurd geonomaet'ormis Martius: a Ma- acknowledge Ms. Betty Loraamm for layan undergrowth palmlet. Principes 16: photographic assistance. 3-10. Kurrn, C. S. 1841. EnumeratioPlantarum. Lrrnutunn Crtnn Stuttgart. 3. MrNlrv, W. D. 7967. Experience with Ecological lif" "y.1" Bllltsten, B. A. 19?0. hardy Ealms in Georgia. Principes 11: ol Euterpe globosa Gaertn. In H. T- 7B-86. Odum and R. T, Pigeon, eds. A tropical Menrrus, C. F. P. vorv. 1838. Historia Na' rain forest, USAEC Technical In{orma' turalis Palmarum. Leipzig. 3: 250. tion Center, Oak Ridge. 8299-8314. Mrrcnulr-, R. S. 1963. Phytogeographyand B.lnnv, D. 1961. Notes on the hardiness of floristic survey of a relic area in the palms on the French Riviera. Principes Marianna lowlands, Florida. Amer. Midl. 5:100-103. Nat. 69: 32V366. Bnowrv,K. E. 1976. Ecologicalstudies of Moonn, H. E. 1963. An annotated check- the cabbage palm, Sabal palmetto. Prin' list of cultivated palms. Principes 7: cipes20: 3-10. 119-182. Deulcnem, B. E. 1936. Index of American L973. The major grouPs of Palms palms. Field Mus. Nat. Hist., Bot. Ser. and their distribution. Gentes Herb. 14: I-456. Il(2): 27-I4L Erlrorr, S. 1817. A Sketch of the Botany 1975' The identity oI RhaPis arun' of South Carolina and Georgia. Charles' d,inacea.Principes 19: 151. I (2). ton. Porrxon, J. 1973. The hardiest palms. Observationson pollina- Essrc, F. B. I97I. Fairchild Trop. Gard. Bull. 2B(2): 11-14. tion in Bactris. Principes \5: 20-24. Sr-*ov, S. G. 1963. Introduction of palms 1973. Pollination in some New in the U.S.S.R. Principes 7: BB-99. Guinea palms. Principes I7: 75-83' Scuurn, R. 1970. Notes on the reproductive Frsuen, J, B. ervn P. B. Toulrrsol. 1973. of Asterogynemartiana (Palmae). Branch and inflorescence production in biology II. Pollination by syrphid flies. Prin' saw palmetto-l (Serenoa repens). Prin- cipes 14: 39-49. cipes 17: 10-19. palm-Rfta' Gr.essultt, S. F. 1972. A revision of B. E' Sn.rr-1, J. K. 1923. The needle Bot. Dahlgren's index o{ American palms. pidophyllutn hystix. J. New York Phanerog. Monog. 6. Gard. 24: 105-114. Henrun, R. M. 1906. Notes on the distri Tonrrrson, P. B. L962. Essays on the bution of some Alabama plants. Bull. morphology of palms VII. A digression Torrey Bot. Club 33: 523-536. about spines. PrinciPes 6: 44-52.