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    reviews TREE vol 1, no 6, December 1988 Population Biology, Social Behavior and Communication in Whales and oo Dolphins Soreness uel aeaict ate often less defined than those of baleen whales. The mating and Peter Tyack calving seasons of odontocetes may not overlap, for the gestation “The baleen whales die from the tothed near the tropics to their summer periods often exceed one year ihales and dolphins ix Me history and in feeding grounds in the Antarctic. Their mating seasons are not as social organization. Even though they During the summer feeding season, synchronized with the annual cycle ‘grow oa larger se, young balen whales adult whales are usually caught as. those of baleen whales, and tend to develop more rapidly than dolpins with full stomachs, while during there may be several. seasons and twothed whales. Except for the the winter breeding season their per year within one odontocete mother-call bond, most grows of Balen stomachs are often empty’. Essen- population. The gestation periods ‘whales are shor-ived, lasting only for tally, baleen whales feast during of the odontocetes tend to increase hous, and indvidual-specic associations the Summer and fast for the rest of with body size (Table 1), and the appear tobe exceptions to te norm. Most the Yeat. larger odontocetes gestate for toothed whales lve in more strucured Birth in most baleen whale spe- about 15 months. By contrast, the groups, in which young animals fave a cies is also highly seasonal: the annual cycle of the baleen whales long period of dependency and socal calves are bom in tropical waters appears to have constrained both learning The communication signals de- during the winter. The gestation the gestation petiod and period of scribed for diferent celacean species have period of two typical baleen whales lactation to under one year. functions suited to the interactions that is nearly one year (Table 1): both Most adontocetes also take lon- predominate in their seis mating and calving occur during the ger to wean their young than ba- same season. Balen whales prob- een whales. Dolphins only 2-3 min ably grow faster than any other length (eg. Tursiops truncatus) may ‘animal. While nursing, 2 calf blue suckle their calves for 18-20 Whales, dolphins and porpoises whale (Belaenoptere musculus) months. The mean duration of suck- all belong to the taxonomic order grows approximately 80 ke/day and ling is 2 years for sperm whales Cetacea. In common English usage, 33 ci/day'. The period of lacta- (Physeter catodon? and 4-5 years all cetaceans more than several tion is relatively short, the calf for the pilot whale species Glo- meters in length are called ‘whales’. being weaned within six months to bicephafamacrorhynchus*. But But the toothed whales such as the one year. AS Table | shows, blue some young of both these two spe- sperm, killer and pilot whales are whales wean at seven months of cies may suckle for much longer much more closely related to dol- age, when the calf is on average periods, Male sperm” whales as phins and porpoises than to the 128meters long? Theirgrowth isso old as 13 years and females up to baleen whales, Dolphins, porpoises rapid that after the short period of 75 years have lactose in theit ‘and toothed whales are all clas- lactation, a calf will be 070% of ts stomachs, indicating the presence Sified together as odontocetes. As length at sexual maturity. On the of milk”. The maximum duration of the name indicates, all odontocetes breeding grounds of humpback suckling has also been estimated have teeth. Balen whales do not whales (Megaptera novaeangliae) for G. macrorhynchus in Japanese have teeth, but instead use baleen one can occasionally identify year- waters where entire schools are to filter small prey from seawater. lings. but animals older than one sometimes caught: comparisons of Baleen whales also have two exter- year can seldom be readily distin- the number of lactating females nal blowholes while odontocetes guished from adults. with the ages of immature animals have only one. Beyond these The usual range of age at sexual in these schools indicate that indi- physical differences, recent studies maturity for baleen whales is 4-10 viduals as old as 15.9 years may still indicate striking differences be: years, with blue and humpback be suckling* ‘ween the two groups in life history, whales maturing as early at 4-5 These extremely long times to social organization and communica: years’. Females of some species weaning have been @ puzzle for tion are capable of having one calf per biologists. Both P. cafodon and CG. year, but a to- or three-year macrorhynchus. start taking. solid Reproductonandfeedingintaleentales breeding cycle is more common, food by the end of ther fist year. Isstrongly seasonal perhaps allowing a female to build The large difference between the ‘Most baleen whales exploit up her fat reserves during feeding mean and maximum ages for wean- schooling planktonic crustaceans or Seasons when she is not pregnant. ing in these species suggests that ‘small fish which abound during the young do not require milk for as summer in polar waters. Each Tong as some of them suckle. It has spring, most species of baleen Yourgtoothed whalesaredependentfor been suggested™ that lactation is whales in the southem hemisphere many years ‘nutritionally important only in the migrate from breeding grounds "Few odontocetes are known to fist few years of life for these spe- have long yearly migrations be- cies. This conclusion is supported ete ack atthe Woods Hole Oearogaphe tween separate feeding and breed- by comparisons with 7. truncatus; Inston, Woods Hole MA C250, USA Ths ing grounds. Nor do they have the although these dolphins wean by atl catia No, 6 9f WHOL annual cycle of feast and famine 1820 months, they stay with the a (9 1, Euever Somos snr 3, Art cies 4880259 TREE vol. 1, no. 6, December 1986 ‘Table hstor parameter for selected cetacean species* Species Gestation lntercalf Age atweaning Lengtnat Sex Ageat Wax Lengthat Max period interval weaning Sovusl longevity? “sexual length® mol ci. maturity jy) fy) maturity (m)_ (nh aleen whales Blue whale? n 2 m0 ve oM 5 26 Boleenoptere musculus F 5 24 A Humpback whale?* now Mime aa Mas 1s ‘Megaptera noveeenlise Fe 2 6 Odontocetes Sperm whale” 1816 58 meanzye 67 M2025 2 3 1% Physatereatodon ax i3 yr F 3 oa aller whale 1% 38) 0 Rm M8 3 58 Orcinus oreo an) ge eal Shor finned plot whale? 15 22 memtsye 27 M17 42st Globicephaig max 13-15 Yr F 9 6 32 45 ‘macrorhynchus Botvenosed dolphin ” 23 18:20m0 ena % 25 38 Tursiope tencatus ew 3238 Harbor porpoiee™ fears smo 1M 46 asa 1s Phosoonaphocoena cee aaa W La Pata river doiphin® ms 2 2-9mo oom ote al 18 Pontopora blames F 27a « Baleon whales are not sizonaly sex have only ben indiosted for femele Deleon whales * The values for any given porametar in tie table were not all derived using the same technique, © Ageing techniques are not as ralabe fot Bsleen wheles as for toothed whales, 60 longevity isnot incited for these species. ty eimorphi,atthough females tena ta grow sight larger than males; maximum length figures mother for the next two years’.The In keeping with the apparent im- males to females is 1.44'°. While prolonged lactation may be seen as portance of social learning, dol- male sperm whales may produce a ritualized display reinforcing the phins are highly skilled at vocal as spermatozoa when as small as 10 m mother-calf bond. However, the well as postural or motor mimicry*. long, the large males over 14 m long. apparent tendency for males to The importance of a jong period (and more than 20-29 years old) are ‘suckle for longer in both species of parental care is marked by a probably the successful breeders* suggests another interpretation. clear postreproductive phase In the Female sperm whales reach sexual Both of these species appear to be life cycle of G. macrorhynchus’. maturity at an average age of 9-10 polyeynous, and prolonged suck- Over 25% of the mature females years’. G. macrorhynchus shows a ling of male offspring may enhance caught their growth and increase the likeli hood of their becoming successful similar to those of postmenopausal males at about 174 breeders, One must be cautious in women), and 25% of these post- interpreting this phenomenon, reproductive females are lactating, Social Interactions among baleen whales however, because in neither spe- Female G. macrorhynchus cease to arebriet cies is it known whether lactating ovulate before the age of 40, but In baleen whales, the most stable females only suckle their own females live for up to 63 years, The bond is between a female and her postreproductive females have calf, and this lasts tess than one Brodie® was one of the first to presumably switched their repro- year. In keeping with their exploita- suggest that the prolonged period ductive effort from producing new ton of patches of prey that are of lactation In odontocetes may be offspring to lactation and caring for highly variable in time and space, related to the importance of social existing offspring, although it is not the groupings of most_ baleen learning. These animals do appear known whether these postrepro- whales during the feeding season to tequire a long period to leam ductive lactating females suckle are temporary. offspring, about feeding, predator avoidance offspring other than their own. the lapanese fishery have similar pattern; the females reach senescent ovaries [histologically sexual maturity at about 9 years, Group structure has been most ‘and social behavior from their — Many odontocete species take intensively studied in humpback ‘mother and other group members. ionger to reach sexual maturity than whales. Humpback whales off New- Young captive dolphins leam many baleen whales, and there tends to foundland feed in groups that are new behavior patterns through it i- be a larger gap between the sexes seldom stable for more than a few tation of adults, For example, cap- in age of onset of sexual maturity. hours; the size of these feeding tive dolphins that were caught in The most extreme example is the groups is often proportional to the the wild are skilled at preparing highly polygynous sperm whale, size of the patches of prey", and large fish for eating, while captive- This species is the most sexually individuals are not identified to- bom dolphins are clumsy in their dimorphic of all cetaceans; the gether in different groups more attempts to imitate this behavior’, length ratio of physically mature often than would be predicted by a 45 TREE vol. 1, no. 8, Decernber 1986 46 = Top to bottom: blue whale, humpback whale, sperm whale, ale ae In the fall, hhurmpack wha- les migrate to winter breed. ing areas near Islands in trop- [eal waters Males have sev- eral strategies for galning access to fe- males on the breed- Ing grounds. Some- times, males join in large groips to fight for access t0 one central random* model. There are a few reports: ffom other areas of more. stable groups of humpback whales, In which individuals appear to learn to coordinate feeding; the same whales may be sighted together. in these groups in, several different feeding’ seasons'%, but their pat- terns of association: have not been subjected to statistical analysis. ‘The traditional: view of whale Biologists has been: that baleen whales are monogamous?, but there is very clear evidence that this is not always the case. In fact, all of the species whose breeding be- havior has’ been’ studied are polygamous. These species, which include humpback, right and. gray whales, favor coastal or shoal wa- ters during’the breeding season (hich is why these species are the most: extensively studied). Again, humpback whales can be used as an example, female", when alone, imales “may instead produce long compll- cated displays, called songs. The song consists of a series of notes and lasts up to 20 minutes before repeating, Since humpback song is sung by males primarily during the breeding season. it has often. been imerpreted asa reproductive advertisement display. Some behavioral observations of Singing whales support this idea. Singing whales tend. to approach other whales nearby, and. when singers join with other whales, they stop singing" If they join with another male, an aggressive interaction usually follows. On the few. occa- sions when. singers have been observed to join with females, behavior associated. with sexual activity is observed. But the pairing Is brief, and whether a male has joined witha female ater singing or as a result-of fighting In a larger group, he only escorts the female fora few hours, When the same females are reidentified, they are seldom seen with-the same male, even after intervals of only several Gays. The same Is true of humpback {groups in general. No groups which Were resighted on the Havatian breeding ground on different days had the same composition’? ‘Odontocetes have stable associations Groups of odontocetes are more structured than those of baleen whales. The members of an odonto- cete group tend to surface with more synchrony than Is typical of baleen whales and the spatial structure of odontocete groups also tends to be more distinct. Balen whales are often spread out in dif- fase herds, and It can be difficult for the observer to ascertain where ‘one group begins and another ends. While individuals of some ‘odontocete species such as the sperm whale may separate by many hundreds of meters when di- ving, members of a group usually regroup to surface at the same time within tens of meters of one another'®. Groups of adontocetes are also much more stable over time than those of baleen whales. The most stable of odontocete groups occur. amongst the killer whales (Orcinus orca) which have been studied ear Vancouver Island, British Columbia. Indivic uals of this species have been {identified in about 30 groups: for over 13 years. These groups seem to be made up of related indi uals: with low birth and death rates, group compasition often does not change for several years at a time. Even though different groups may travel together for a few weeks, no permanent exchange of any anirial between groups hasbeen observed"? No other odontocete groups are known to be as stable as those of the killer whales. There are gener- ally fewer mature males than fe- males in groups with young, and ‘males do not tend to remain in the ‘same group as long as females do. TREE vol. 1, no. 6 December 1986 In many odontocete species, adult males may associate with female ‘groups for only a few days at a time, ‘As young males mature, they tend to leave thelr natal group and form juvenile groups, which may also contain juvenile females in some species. The long mother-calf bond provides the stablest unit of group structure In odontocete species. Odontocete groups are typically formed of females with thelr your sometimes spanning several ge: erations. Mark-recapture studies of sperm whales have shown that mi ture females tagged within a group are likely to be caught from the same group even after Intervals of 5-10 years"®. This is not the case with mature - males. Studies of bottlenosed dolphins inhabiting shore waters show that group com- position in. this species changes frequently"®, However, clusters of females that share preferred habi- ‘tats tend to be sighted in the same ‘groups, Males tend to range over a Wider area and may even leave the well defined population bound- aties for periods of several month’ ‘Animals -- within. odontocete groups cooperate in. a. variety of ‘ways; in particular, allomaternal be- havior is commonly reported. In captive, groups of dolphins, non- pregnant females often closely attend the birth of a calf and have been observed to break, the, umbi- Jical cord. These ‘aunts'-swim near the mother-calf. pair, and: young bottlenosed dolphin’ calves will swim with one of these aunts while the mother is feeding. The appar- ent importance of allomatemal be- havior in odontocetes -stands in Solpnia strong contrast to the behavior of humpback whales; when female humpback whales have. a” young calf, they avoid other whales™, Care-giving behavior has been reported surprisingly’ often ~ for codontocetes, ever’ between mem- bers of different species. Wild and captive odontocetes will stand by, assist, or physically support dis- tressed or. Injured individuals” ‘Typically, a’pair of healthy animals will move to either side of 2 dis- tressed one andiftit up to breathe in a highly coordinated fashion, ‘Social defense from predators is. also considered to be a major func- tion of odontocete schools, and td bbe a determinant of their size and structure. The responses to disturb- lance of spotted dolphins (Stenella attenuata) in the Pacific are among the best studied”. Upon disturb- ance, these dolphins form a tight school. Mothers with young raft in the center of the group and-are protected by other adults diving continuously under the raft and sur- facing on al sides of it ‘Cooperative behavior in edonto- cetes also extends to feeding. Food sharing has been reported for several species’. Many odonto- cetes have highly coordinated pat- tems of searching for prey, and Botton fo tp. shor fred pllot whale. boule: nosed dolphin, harbor porpoise, Lt Pata river there is evidence that Individuals may display to advertise the loca- tion of prey®. Many. odontocete species herd fish in order t0 cap- ‘ture prey’; during. this process, some individuals that are herding fish appear to forego feeding while other individuals feed. The struc- ture of odontocete groups and the manifold reports of cooperation jn these animals have led to specula- tion that odontocetes maintain individual-specific social relation- ships through reciprocal altruism”. Large assemblages of -odonto- eetes. are typically. made up. of smaller subgroups: that are often distinctly separated from one another and which remain together even after the larger assemblages break up. However, in some spe- cles, includling the bottlenosed dot- pphin, interchange of animals. be- tween subgroups within a larger -_ — SPRAY ScorTy 15 = 15 z 2 2 Pe 7 ‘ es 3 = : = % Os 0 0.5 10 MK a5 SCOTTY 5 = SPRAY om 151 SE 1 =z © 104% 10) Wf 5) fe eS AES 0. o 04 08 Fig 1. Sound spactograms of whistles recorded from two captive betlenosed dalphins (Turis frais a forale named Spray and @ male named Seoty On the Tet of the top row isthe signature thse of Spray on the top right isan ination of thls whlstle produced by Sezty On the Bottom lft the signature whist of Scary on the Botom right an ination produced by Spry. The horizontal [less time in seconds, Repu with ermision, from R28. assemblage is common". Never- theless, individuals may show strong patterns of association with other individuals, even within otherwise fluid pattems of group- ing. For example, some pairs of adult male bottlenosed dolphins are almost always sighted together, even jn different groups, for many years’. As odontocetes’ develop, they may leave their natal group and join different groups for long. Periods. The combination of highly structured pattems of association Frequency (kHz) Time (sec) Fig.2. Sound spectrograms of codas produced by tro spem whales (Prise leon) as they approached one hotter Such codes typeally lifer i both *ythme pattem and number of clicks (the coda above fas Seven clicks, that below has rine) Rewaducl oie pert, fom Rel 28 48 between individuals coupled with ‘occasionally fluid patterns of social grouping argues that individual- specific social relationships are im- portant elements of the social structure of many odontocete spe- cies. What we know about com- munication in dolphins and sperm whales reinforces this view. Individual distinctive signals in dolphins and sperm whales Dolphins of many species pro- duce high-pitched pure tone whis- tle sounds which are modulated in frequency. Recordings from over 100 captive dolphins have shown that each adult dolphin, when iso- lated, tends to produce one certain kind of whistle which is distinct from the whistles of other dolphins in its group”. These whistles have been called’ signature whistles: they apparently function to broad- cast the individual identity of the whistler. Bottlenosed dolphin calves produce unstereotyped whistles on their first day of life; most develop a stereotyped whis- tle within their first year. While dol- pins are interacting, they tend to produce their own signature whi tle, but they do also mimic the signature whistles of others in their tank (Fig. 1), What might the func- tion of this mimicry be? Carefully controlled experiments have shown that dolphins are skilled at mimick- ing man-made whistle-like sounds, and that dolphins can be trained to label arbitrary objects with these arbitrary whistles®”. Perhaps such cognitive skills function in un- trained dolphins to allow them to label other members oftheir social TREE vol. 1, no. 6, December 1986 groups by mimicry of their signa. ture whistles, Sperm whales also produce a sound that appears to be indi- vidually distinctive, with an acous- tic structure completely different from dolphin whistles; the sound takes the form of a short series of clicks, called codas, with highly stereotyped chythmic patterns (Fig. 2}, Several lines of evidence indi- cate that each whale within a group has a distinctive coda. When the Tocations of whales producing codas are calculated, similar codas appear to come from the same source, while different codas come from different sources. The sound Tevel and timbre of the clicks that make up codas also tend to be similar for the same codas and different for other codas during re- cording periods of an hour or so. Like dolphins and their signature whistles, sperm whales can mimic the codas more typical of other individuals In well defined coda exchanges* (Communication in other cetacean species Individually distinctive signals have not been definitely identified for any of the baleen whales nor for several groups of odontocetes. Not enough is known about social com- ‘munication in any of these species for one to conclude categorically that they do not use vocalizations 10 communicate individual identity. ‘More extensive studies may reveal such signals in the future. However, ‘most of these species have social structures that differ from the whis- ting dolphins and the coda- producing sperm whales. The sig nals that have been described for these species have functions that appear more suited to the kinds of interaction that predominate in their societies. Some dolphin species, including the platanistid river dolphins and phocoenid porpoises, have never een reported to whistle®. Inter- estingly, these animals are more similar to baleen whales than other dolphins in several respects”. As the last two rows of Table 1 show, their timespans for gestation, wean- ing and sexual maturation are simi lar to those of baleen whales. Little ig known about the social structure of these dolphins, but many sight ings are of lone animals; when not TREE vol. 1, no. 6, December 1986 alone, they tend to be in groups of only a few animals, They are clearly ‘much less social than the dolphin species that do whistle. There are no reports of indi- Vidually distinctive signals for killer whales, but they do produce group- specific dialects for one sound — the 'S' call, Each killer whale ‘group has a different repertoire of S alls, and apparently every indi- vidual within the group produces each $ call, As was mentioned above, even when different groups ‘mingle, they always separate into the original groups. Whereas dol- phins and sperm whales apparently use their individually distinctive signals to maintain associations be- tween individuals within changing ‘groups, the communication system described for killer whales appears to function to maintain the cohes- Iveness of their stable groups. Little is known about the social function of sounds produced by baleen whales during the feeding season, but there are some hints that humpback wales and finback whales (Balaenoptera physalus) may produce individually distinctive sounds while feeding in groups. ‘This may help to maintain the rare associations of individuals that feed together In a coordinated fashion. During the breeding season, several species of baleen whales produce advertisement displays The best studied of these are the songs of the humpback whale. Not nly has humpback song not been shown to be Individually distinc- tive, but ithas an acoustic structure that renders individual identifica- tion extremely difficult. Humpback songs recorded within a few weeks of each other on the same breeding ‘ground are very similar. But every aspect of the songs changes gradu. ally throughout each singing season”, Sounds may change in pitch, duration and timbre; they may disappear from the song en- tirely, and new sounds may appear in some other part of the song. These changes are cumulative. Over a twenty-year period, entire songs are transformed: not one sound from the song's first version Is left intact in the last™® As Fig 3 indicates, whales singing at any one time within a population sing very imilar sounds compared t0 the anges that occur within even one ‘Young humpback whale off Hawai Pl by Grane Es year", This Is very different from the signature whistles of dolphins, which are stable over periods of many years Future research Future studies should reveal whether these associations be- tween social organization and com- ‘munication reflect fundamental dif. ferences in the social relations of different cetacean groups. Long term studies of the social behavior and pattems of association be- tween identified individuals are under way for many cetacean spe- cies. Current studies of social com- ‘munication in cetaceans are also advancing rapidly. New techniques have been developed to locate sound sources underwater and to identify which animal produces which call. These will facilitate study of the social functions of ceta- cean signals and in particular, should help to determine whether other cetacean species produce in- dividually specific signals. References 1 Lockyer ¢. (19811 FAO. Fk Sr 5 (3, 2 ck, (1984 Ren Wha Comm Sp lua 67755. 5 Sest.PB.Cenham, PAS and MacLeod, N ‘9841 epi Wha Come Spec ue 651279 4 guy, and Marsh (1988 Rep et ‘wha Come, Spe tu 6, 259-310 Wells RS. Seat. MD. andivine, AB. Cur anal in press) (Bogle, PF {1963 An. Mi Nat 82 size 1 Nom, KS, and Doh (1960) in Cte Beker Meck ad Encons (Herman, {Entved pp. 211-21, Wiley Iterscence 1 Herman (1960 Cane Bea Mecano nd Fuacons (Herman, LM. bp. 365-129 Wiley Inercience S hiarah Wand Kasoy T1968) Rep et ‘Wha Come Se sue 6311-335 10 Best P1979 Serra Marne or WiisO fe et ae Spel a Unidgtited ) /' | Fig. 3, Sound spectrogeains of ane phase trom the songs of hamphack whales Mapes woaeangie) retoried In Hawallan waters. The phrases onthe fp row were sung bythe ame individual, labeled vs approximately one year apart Te phrases onthe lowe ow weve corded fom unidertNed ‘whales (very likely to be diferent ndsiduals) wtn two weeks ofthe phrases directly above them on the top row It is obvious thatthe pases of diferent whales sung within the same to week interval vera comparsos) ore auch more similar han phases the sere ndvidual whale wth one Yea? Inter top horzanalcomparcon| Repofuc, ik poms, om a 34 “9 Regeneration of Canopy Trees in inl (Cao ests it Research, Vl 3) tenn (Winn, HE and Ola BL ed, pp. 1212289 Plenum Press TH Whitehead, Harcourt, Ingham. K find Cle, (0981 Cae 2 58 687-92 {2 Baker, CS. and Herman, UM, i885) at His 9452-6 13 Tyack, Pend Whitehead 1983) eda, 152-154 [eTyace P1981) Bhs Fal Soil los-ti6 15 Mobley, JR. Jrand Hemman, LM (1985 on | dal 3, 02-172 fs Watkne, WA. and Schevil E1977) DowpSee Riseth 26, 693-698 1 Bigg MA (19821 Rept Wha Comm, 32, esate Is OheumsS. (9711 el Rep. Whales Res. st Bias shane, SH, Wells RS. and Wars B (i986) Mer Mana Se 234-63 20 Herman LM and Tavolga WN. (1980) Tropical Wet Forests Deborah A. Clark 180 ‘The most diverse tre communities on earth, the topical wet forests, 10-@ large degree remain ecological enigmas. What ‘accounts forthe coexistence of 100 oF more tree species per Reclare, compared to the 15. or fewer found in mest temperate forests? What are the Hfespans of topical forest tees? What factors control their populations through time and space? Do the different species have kighly individual regeneration paterns, or are many in fact ecological equivalents? Although we are far from having satslcory answers to these question, recent studies of regenera tion praceses'are leading toward new interpretations ofthese complex commun tis. Until recently, surprisingly litle detail was available about the pro- cess of regeneration of trees in tropical forests, Whereas the annual rings formed in the wood of temperate trees can reveal each individual's age and growth history, this fundamental tool of the tree population biologist cannot be used with most tropical species be- cause they do not form clear annual Debowh Ct Isat La Seka Bilgkal Staton ‘Organization fo Tec Stuties, Apaade 676 250 San Peo, Cosa ic, econ Basi, Mechanin ond Functor (Hearn Ue) pp. 49-209, Wey Intercence 21 caldvell MC and Cale, DK (196) in ‘Whak, Dans, and Pes (Noms, KS, fed, pp. 755-189, Univesty 9f califomis Frese 2 Connor, RC. and Nor, KS. (1981) Am Nat 119, 356-574 2B Wess, 5.1986) In Dep Coon ad Belavir AConpoate Apa Schusterman, Ri. Thomas IA. nd Wood, FG. eds), 347-350, Enbaim Ascocates Maral 1.219-260 2 Cale N.C end Caldwell DK. (1979 in Behar of Mame imal Care Pesce Resa, Vl 3) Cea (Winn HE and Oils: BL ede, pp 369-10, Plenum Press 2 Tyack P1986) Bes Ee Sel 18 Bis? 2 Aichards, 0G. Wot LP and Herman. LEM 119841T Com Pst 96, 10-28 growth rings'. A second major chal- Tenge for the study of tropical tree ecology Is presented by the spe- ies diversity itself; in relatively few sites have all the tree species been identified. Furthermore, most tree species In tropical forests occur at densities of

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