Animal Diversity Web

Ge­o­graphic Range

Ha­pale­mur au­reus, the golden bam­boo lemur, is en­demic to the trop­i­cal, moist, south­east­ern rain­forests of Mada­gas­car. It was first ob­served liv­ing sym­patri­cally with other lemur species in 1985. Since its dis­cov­ery, fur­ther stud­ies have con­firmed the range of H. au­reus over­laps with Ha­pale­mur griseus and Pro­le­mur simus, mostly re­stricted to the south cen­tral part of the east­ern forests. Sev­eral groups live in and around Ra­nomafana Na­tional Park in Mada­gas­car, a park that cov­ers ap­prox­i­mately 43,500 square hectares of rain­for­est. Ha­pale­mur au­reus does not seem to be re­stricted by al­ti­tude, as groups have been found through­out an al­ti­tude range of 800 to 1300 m within Ra­nomanafana Na­tional Park. In 1993, H. au­reus was also found in An­drin­gi­tra Na­tional Park, south of Ra­nomafana. These two areas are con­nected by a for­est cor­ri­dor, which is be­com­ing in­creas­ingly frag­mented. One sur­vey dis­cov­ered Ha­pale­mur au­reus in 4 out of 8 sites along this cor­ri­dor. There have not been sight­ings north of Ra­nomafana or south of An­drin­gi­tra and a ten­ta­tive south­ern limit is placed at the Man­am­pa­trana River. (Ar­rigo-Nel­son and Wright, 2003; Good­man, et al., 2001; Irwin, et al., 2005; Meier, et al., 1987; Mutschler and Tan, 2003; Tan, 1999)

• Biogeographic Regions
• ethiopian
  • native

• Other Geographic Terms
• island endemic

Habi­tat

Golden bam­boo lemurs are ar­bo­real pri­mates re­stricted to bam­boo patches in the south­east­ern Mada­gas­car rain­for­est. With a diet spe­cial­iz­ing in giant bam­boo (Cathar­iostachys mada­gas­caren­sis), it is not sur­pris­ing to find bam­boo as an im­por­tant habi­tat re­quire­ment for golden bam­boo lemurs. A 2003 sur­vey of lemurs in Ra­nomafana Na­tional Park doc­u­mented a min­i­mum of two C. mada­gas­caren­sis stands per site. Each bam­boo stand av­er­aged 250 by 200 m in four sep­a­rate sites, and groups golden bam­boo lemurs were found liv­ing near each one. In the same sur­vey, two other sites with ev­i­dence of golden bam­boo lemur ac­tiv­ity that did not con­tain C. mada­gas­caren­sis, had a dif­fer­ent species of large-culm bam­boo, Arun­d­i­naria am­bositren­sis. Other species of bam­boo grass and small-culm bam­boo (genus Cephalostachyum) were also found at each site. All types of bam­boo species can co-ex­ist within the same area, but small-culm bam­boo and grasses were found with the great­est abun­dance in edge for­est habi­tats and in gaps in the for­est cre­ated by tree falls. The suc­cess­ful and rapid col­o­niza­tion of bam­boo in dis­tur­bance areas can often cre­ate an over­lap of H. au­reus habi­tat with human ac­tiv­i­ties such as log­ging and agri­cul­ture. If these lemurs live near a for­est edge habi­tat, new moth­ers tend to move into more shel­tered, dense, habi­tats for the first 10 to 14 days after giv­ing birth. (Ar­rigo-Nel­son and Wright, 2003; Mutschler and Tan, 2003; Tan, 1999)

• Habitat Regions
• temperate
• tropical
• terrestrial

• Terrestrial Biomes
• rainforest

• Range elevation

  800 to 1300 m

  2624.67 to 4265.09 ft

Phys­i­cal De­scrip­tion

Bam­boo lemur species gen­er­ally have round heads, large faces, short muz­zles, and small ears hid­den by fur. They pre­fer ver­ti­cal rest­ing pos­tures and their tails are not pre­hen­sile. The hands and feet are short and broad with large pads under the tips of toes and fin­gers. All teeth have a ser­rated cut­ting edge, ex­cept for the mo­lars. These cut­ting edges are thought to be an adap­ta­tion for a coarse bam­boo diet. The spe­cific den­tal for­mula for this species has not been recorded; how­ever, the den­tal for­mula in the fam­ily Lemuri­dae is I 2/2, C 1/1, PM 3/3, and M 3/3 = 36. As strep­sirhines, lemurs pos­sess a den­tal comb or tooth­comb, a for­ward pro­jec­tion of the lower jaw teeth often used in groom­ing. Den­tal pe­cu­liar­i­ties exist among the pre­mo­lars of bam­boo lemur species, most likely due to their bam­boo rich diets. Bam­boo lemurs have ad­di­tional cusps on the upper fourth pre­mo­lars. These cusps are thought to help in crush­ing the fi­brous plant ma­te­r­ial that they eat. Golden bam­boo lemurs can eas­ily be dis­tin­guished from other bam­boo lemur species by the pale golden fur on their bel­lies, inner limbs, cheeks, throat, and sur­round­ing the eyes. Other bam­boo lemur species have fur col­ors rang­ing from sooty grey coats to rusty brown, with the oc­ca­sional “oli­va­ceous” hue in Ha­pale­mur griseus. The ears of golden bam­boo lemurs have golden-brown tips and are not tufted. The coat is dor­sally red­dish-brown; darker over the shoul­ders, back, top of head, and tail. The tail has a dark­ened tip. Five bam­boo lemur species are rec­og­nized. Golden bam­boo lemurs are the largest, with an adult weight of 1.25 to 1.7 kg. Head-body length is ap­prox­i­mately 34 cm, while tail length is about 41 cm, for a total length of 70 to 80 cm. Fe­male and male golden bam­boo lemurs are sex­u­ally monomor­phic. Fe­males have only one pair of mam­mae, lo­cated pec­torally. They have a diploid com­ple­ment of 62 chro­mo­somes, all of which are acro­cen­tric. Golden bam­boo lemurs have well-de­vel­oped brachial glands on the inner upper arm, and an el­lip­ti­cal hair­less patch on the lower arm, pos­si­bly with a carpal gland. Elbow flex­ion brings the two glands to­gether. The brachial gland se­cretes a white, sticky fluid with a strong smell. (Jern­vall, et al., 2008; Meier, et al., 1987; Mit­ter­meier, et al., 2006; Mutschler and Tan, 2003; Nowak, 1999; Ya­mashita, et al., 2009)

• Other Physical Features
• endothermic
• homoiothermic
• bilateral symmetry

• Sexual Dimorphism
• sexes alike

• Range mass

  1.25 to 1.7 kg

  2.75 to 3.74 lb

• Average length

  75 cm

  29.53 in

Re­pro­duc­tion

Golden bam­boo lemurs are monog­a­mous and mat­ing takes place in July or Au­gust. There is some ev­i­dence that male testis size in­creases prior to the breed­ing sea­son. One study showed a male, cap­tured in May, hav­ing a tes­tic­u­lar vol­ume of 17.414 cm3. The same male, cap­tured again in Oc­to­ber, had a tes­tic­u­lar vol­ume of 6.351 cm3. Fur­ther stud­ies are needed to in­ves­ti­gate this find­ing. (Glan­der, et al., 1992; Norosoari­naivo and Tan, 1998; Tan, 1999b; Ya­mashita, et al., 2009)

• Mating System
• monogamous

Ges­ta­tion lasts ap­prox­i­mately 138 days, and golden bam­boo lemurs give birth just once per year, usu­ally to a sin­gle off­spring. Al­tri­cial young are born in De­cem­ber and re­quire sev­eral months of ma­ter­nal care and lac­ta­tion. Un­like other bam­boo lemur species, fe­males nest with their off­spring for the first 10 to 14 days in se­cluded areas of dense veg­e­ta­tion. Wean­ing takes place when off­spring are 6 to 8 months old. (Glan­der, et al., 1992; Tan, 1999b)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• viviparous

• Breeding interval

  Golden bamboo lemurs breed once yearly.

• Breeding season

  Golden bamboo lemurs breed from July to August.

• Average number of offspring

  1

• Average gestation period

  138 days

• Range weaning age

  6 to 8 months

• Average time to independence

  3 years

Golden bam­boo lemurs prac­tice in­fant park­ing and oral trans­port. Moth­ers leave in­fants while for­ag­ing and can travel as far as 250 me­ters away. The in­fant is left for an av­er­age time of 200 min­utes. There are no pub­lished ob­ser­va­tions of pa­ter­nal care, but it is a pos­si­bil­ity con­sid­er­ing the monog­a­mous be­hav­ior of this species. Sib­lings have also been ob­served to care for in­fants. Off­spring live with their par­ents as a fam­ily group until ap­prox­i­mately 3 years of age, after which they dis­perse. (Norosoari­naivo and Tan, 1998; Tan, 1999b)

• Parental Investment
• altricial
• female parental care
• pre-fertilization
  • provisioning
  • protecting
    • female
• pre-hatching/birth
  • provisioning
    • female
  • protecting
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female
• extended period of juvenile learning

Lifes­pan/Longevity

Av­er­age lifes­pan of H. au­reus is not yet known. A sis­ter species, Ha­pale­mur griseus, lived in cap­tiv­ity until the age of 23. On av­er­age, H. griseus fe­males and males live to 17.1 years and 12.8 re­spec­tively. (Ha­keem, et al., 1996; Mit­ter­meier, et al., 2006)

Be­hav­ior

Golden bam­boo lemurs live in small groups, not often ex­ceed­ing four in­di­vid­u­als. Usu­ally the group con­sists of an adult male and fe­male, with smaller young adults or ju­ve­niles. The group is a fam­ily, com­prised of a monog­a­mous pair and their off­spring. Vary­ing ac­counts list golden bam­boo lemurs as di­ur­nal or cre­pus­cu­lar, with a dis­tinct mid-day rest pe­riod. The ma­jor­ity of wak­ing hours are spent eat­ing and for­ag­ing. (Gar­butt, 1999; Meier, et al., 1987; Mit­ter­meier, et al., 2006; Tan, 1999)

• Key Behaviors
• arboreal
• scansorial
• diurnal
• crepuscular
• motile
• sedentary
• territorial
• social

Home Range

A study of two sep­a­rate fam­ily groups of golden bam­boo lemurs over a two-year pe­riod showed their mean home range to be 26 hectares. An­other group, an adult pair and two off­spring, had a home range of 80 hectares. Ter­ri­to­ri­al­ity has not been doc­u­mented in Ha­pale­mur au­reus. But the pres­ence of a brachial gland and sub­se­quent re­ports of scent mark­ing in a sis­ter species, Ha­pale­mur griseus griseus, sug­gests they may be ter­ri­to­r­ial. (Tan, 1999; Wright and Ran­d­ri­manan­tena, 1989)

Com­mu­ni­ca­tion and Per­cep­tion

Golden bam­boo lemurs are highly vocal, with at least two dis­tinct calls. They were first recorded as per­form­ing a “hard grunt” and a loud call at night. One call is de­scribed as a quiet “res­o­nant wuulp with in­quis­i­tive in­flec­tion.” This sound pos­si­bly serves as a con­tact call be­tween mem­bers of the fam­ily group, as it is some­times re­sponded to and re­peated by oth­ers. A longer call is de­scribed as a loud, sharp, throaty honk that de­creases in vol­ume as it is re­peated. This longer call ap­pears to be de­liv­ered by only one mem­ber of the group and is often heard along with the "wuulp" call. This sec­ond call may have a ter­ri­to­r­ial func­tion. Golden bam­boo lemurs, like other lemurs, have binoc­u­lar vi­sion. The func­tion of the brachial gland in Ha­pale­mur au­reus is not yet known. Lesser bam­boo lemurs use scent-mark­ing, vocal dis­plays, and chas­ing in ter­ri­tory de­fense. Ring-tailed lemurs par­tic­i­pate in ter­ri­to­ri­al­ity via scent mark­ing. They ex­crete a fatty sub­stance from their brachial glands, which is rubbed onto its long tail for scent dis­per­sal. The ex­cre­tions of the brachial gland in golden bam­boo lemurs are likely used in a sim­i­lar way. (Gar­butt, 1999; Meier, et al., 1987; Mutschler and Tan, 2003; Nowak, 1999)

• Communication Channels
• visual
• acoustic
• chemical

• Other Communication Modes
• scent marks

• Perception Channels
• visual
• tactile
• acoustic
• chemical

Food Habits

Golden bam­boo lemur diet is pri­mar­ily com­posed of Mada­gas­car giant bam­boo, but Cephalostachyum vigu­ieri, Arun­d­i­naria am­bositren­sis, and other bam­boo grasses are also con­sumed. Bam­boo is con­sumed year round, as is the con­sump­tion of small amounts of soil and fungi. Al­though ap­prox­i­mately 78% of golden bam­boo lemur diet is giant bam­boo, this species has been ob­served to eat other bam­boo and grass species (10%), bam­boo fo­liage (3%), fruit (4%), and other (5%). Small in­stances of fru­givory are op­por­tunis­tic. Golden bam­boo lemurs pre­fer to con­sume young growth of giant bam­boo, such as the leaf base, shoots, and pith, dis­card­ing more ma­ture growth. New growth bam­boo has a lower en­ergy cost dur­ing the mas­ti­ca­tion and is more eas­ily di­gested. It also has higher con­cen­tra­tion of pro­tein. Young shoots are only sea­son­ally avail­able. Golden bam­boo lemurs con­sume vary­ing parts of bam­boo when the young shoots are less abun­dant dur­ing dif­fer­ent sea­sons. Pre­vi­ously, it was pro­posed that niche par­ti­tion­ing ex­plained the pref­er­ence of dif­fer­ent bam­boo por­tions in Ha­pale­mur au­reus and two other species of bam­boo lemurs that live sym­patri­cally: Ha­pale­mur griseus and Pro­le­mur simus. These species ap­peared to se­lect al­ter­nate por­tions of bam­boo plants. Sub­se­quent stud­ies cast doubt on the de­gree of niche par­ti­tion­ing after find­ing over­lap in the diets of all three species. Iron­i­cally, the most pro­tein-rich part of bam­boo is also the most deadly; giant bam­boo shoots con­tain high amounts of cyanide. Be­cause golden bam­boo lemurs specif­i­cally tar­get these shoots for con­sump­tion, they are es­ti­mated to in­gest about 500 g of bam­boo daily, which con­tains ap­prox­i­mately 12 times the lethal dose of cyanide in com­pa­ra­ble mam­mals. More re­cent stud­ies sug­gest the cyanide con­cen­tra­tion in Cathar­iostachys mada­gas­carien­sis shoots to be up to four times higher than orig­i­nally es­ti­mated. A pos­i­tive test for cyanide has been found in the urine of H. au­reus, but only rarely in fecal sam­ples. This re­sult sug­gests cyanide is ab­sorbed by the gas­troin­testi­nal tract with the kid­neys in­volved in its ex­cre­tion. It takes more than 20 hours for com­plete di­ges­tion and elim­i­na­tion of fecal mat­ter. The means by which they avoid cyanide poi­son­ing is un­known. (Ball­horn, et al., 2009; Glan­der, et al., 1989; Tan, 1999; Ya­mashita, et al., 2010; Ya­mashita, et al., 2009)

• Primary Diet
• herbivore
  • folivore
  • lignivore

• Plant Foods
• leaves
• wood, bark, or stems
• fruit

Pre­da­tion

Di­rect hunt­ing by hu­mans is the most widely re­ported type of pre­da­tion on golden bam­boo lemurs. Two types of hunt­ing have been re­ported: pro­jec­tile, usu­ally using spears, slings and blow­guns; and trap­ping, in­volv­ing snares. (Gar­butt, 1999; Irwin, et al., 2005; Mit­ter­meier, et al., 2006; Mutschler and Tan, 2003)

• Known Predators

  • humans (Homo sapiens)

Ecosys­tem Roles

The ecosys­tem role of Ha­pale­mur au­reus has not been stud­ied. Fru­giv­o­rous lemurs have been im­pli­cated as im­por­tant vec­tors for seed dis­per­sal in Mala­gasy rain­forests, but it is un­known what, if any, role they may play in prop­a­ga­tion of bam­boo. (Dew and Wright, 1998)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Golden bam­boo lemurs are part of the fas­ci­nat­ing and unique en­demic lemur fauna of Mada­gas­car.

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no ad­verse im­pacts of golden bam­boo lemurs on hu­mans. (Gar­butt, 1999; Mutschler and Tan, 2003)

Con­ser­va­tion Sta­tus

The most re­cent IUCN Red List as­sess­ment (2008) con­cluded that golden bam­boo lemurs are en­dan­gered due to their small range and habi­tat frag­men­ta­tion. The species is very rare, re­stricted to the south­east­ern rain­for­est in Mada­gas­car near areas of bam­boo growth. Re­cent sur­veys have iden­ti­fied two re­gions in which the for­est cor­ri­dor be­tween Ra­nomafana and An­drin­gi­tra have be­come dis­con­tin­u­ous. This cre­ates a bar­rier to mi­gra­tion and gene flow. A 2005 sur­vey es­ti­mated the pop­u­la­tion to be 5,916 in­di­vid­u­als evenly dis­trib­uted through­out the area. Even dis­tri­b­u­tion is likely an un­re­al­is­tic as­sump­tion, so the pop­u­la­tion is es­ti­mated at 25% less than this cal­cu­lated value. Other sources sug­gest the pop­u­la­tion to be as low as 1,000 in­di­vid­u­als. (Gar­butt, 1999; Glan­der, et al., 1989; Irwin, et al., 2005; Suss­man, 2002)

• IUCN Red List

  Endangered
  More information

• IUCN Red List

  Endangered
  More information

• CITES

  Appendix I

Other Com­ments

Golden bam­boo lemurs were first dis­cov­ered in 1985 in Ra­nomafana Na­tional Park in south­east­ern Mada­gas­car and first de­scribed in 1987. (Fausser, et al., 2002; Meier, et al., 1987; Pas­torini, et al., 2002)

Con­trib­u­tors

Dana Kowal­sky (au­thor), Uni­ver­sity of Man­i­toba, Jane Wa­ter­man (ed­i­tor), Uni­ver­sity of Man­i­toba, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Ref­er­ences

Ar­rigo-Nel­son, S., P. Wright. 2003. Sur­vey re­sults from Ra­nomafana Na­tional Park: new ev­i­dence for the ef­fects of habi­tat pref­er­ence and dis­tur­bance of Ha­pale­mur. Folia Pri­ma­tol, 75: 331-334.

Ball­horn, D., S. Kautz, F. Rako­toariv­elo. 2009. Quan­ti­ta­tive vari­abil­ity of cyano­gen­e­sis in Cathar­iostachys mada­gas­carien­sis – the main food plant of bam­boo lemurs in south­east­ern Mada­gas­car. Am J Pri­ma­tol, 71: 305-315.

Dew, L., P. Wright. 1998. Fru­givory and Seed Dis­per­sal by Four Species of Pri­mates in Mada­gas­car's East­ern Rain For­est. Biotrop­ica, 30/3: 425-437.

Fausser, J., P. Pros­per, G. Do­nati, B. Ra­mana­man­jato, Y. Rum­pler. 2002. Phy­lo­ge­netic re­la­tion­ships be­tween Ha­pale­mur species and sub­species based on mi­to­chon­dr­ial DNA se­quences. BMC Evol Biol, 2: 1-9. Ac­cessed No­vem­ber 01, 2013 at http://​www.​biomedcentral.​com/​1471-2148/​2/​4.

Gar­butt, N. 1999. Mam­mals of Mada­gas­car. New Haven and Lon­don: Yale Uni­ver­sity Press.

Glan­der, K., P. Wright, D. Sei­gler, V. Ran­dri­ana­solo, B. Ran­dri­ana­solo. 1989. Con­sump­tion of cyanogenic bam­boo by a newly dis­cov­ered species of bam­boo lemur. Am J Pri­ma­tol, 19: 119-124.

Glan­der, K., P. Wright, P. Daniels, A. Meren­len­der. 1992. Mor­pho­met­rics and tes­ti­cle size of rain for­est lemur species from south­east­ern Mada­gas­car. J Hum Evol, 22: 1-17.

Good­man, S., V. Razafind­rat­sita, H. Schutz, R. Rat­sim­bazafy. 2001. In­ven­taire Bi­ologique du Parc Na­tional de Ra­nomafana et du couloir forestier qui relie au Parc Na­tional d'An­drin­gi­tra. Pp. 231-243 in S Good­man, V Razafind­rat­sita, eds. Les Lemuriens. An­tana­narivo, Mada­gas­car: CIDST.

Ha­keem, A., R. San­doval, M. Jones, J. All­man. 1996. Brain and life span in pri­mates. Pp. 78-104 in J Bir­ren, ed. Hand­book of the Psy­chol­ogy of Aging. Waltham, MA: Aca­d­e­mic Press.

Irwin, M., S. John­son, P. Wright. 2005. The state of lemur con­ser­va­tion in south-east­ern Mada­gas­car: pop­u­la­tion and habi­tat as­sess­ments for di­ur­nal and cath­e­meral lemurs using sur­veys, satel­lite im­agery and GIS. Oryx, 39/2: 204-218.

Jern­vall, J., C. Gilbert, P. Wright. 2008. Pe­cu­liar tooth ho­molo­gies of the greater bam­boo lemur (Pro­le­mus simus). Pp. 335-342 in C Gilbert, J Flea­gle, eds. Elwyn Si­mons: A Search for Ori­gins. He­lin­ski, Fin­land: Springer.

Meier, B., R. Al­bi­gnac, A. Peyieras, Y. Rum­pler, P. Wright. 1987. A new species of Ha­pale­mur pri­mates from south­east Mada­gas­car. Folia Pri­ma­tol, 48: 211-215.

Mit­ter­meier, R., W. Kon­stant, F. Hawkins, E. Louis, O. Lan­grand, J. Rat­sim­bazafy, R. Ra­soloari­son, J. Ganzhorn, S. Ra­jao­belina, I. Tat­ter­sall, D. Mey­ers. 2006. Lemurs of Mada­gas­car. Wash­ing­ton, DC: Con­ser­va­tion In­ter­na­tional.

Mutschler, T., C. Tan. 2003. Ha­pale­mur, bam­boo or gen­tle lemurs. Pp. 1324-1328 in S Good­man, J Ben­stead, eds. The Nat­ural His­tory of Mada­gas­car. Chicago: Uni­ver­sity of Chicago Press.

Norosoari­naivo, J., C. Tan. 1998. In­fant care in Ha­pale­mur au­reus, Ra­nomafana Na­tional Park, Mada­gas­car. Con­gress of the In­ter­na­tional Pri­ma­to­log­i­cal So­ci­ety (ab­stracts), 17: 384.

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Pas­torini, J., M. Forstner, R. Mar­tin. 2002. Phy­lo­ge­netic re­la­tion­ships of gen­tle lemurs (Ha­pale­mur). Evol Pri­ma­tol, 1: 150-154.

Stew­art, I. 1988. Dis­cov­ery of a new species of lemur. Na­ture, 333: 206.

Suss­man, R. 2002. Adap­tive array of lemurs of Mada­gas­car re­vis­ited. Evol An­thro­pol, 1: 75-78.

Tan, C. 1999. Group com­po­si­tion, home range size, and diet of three sym­patric bam­boo lemur species (genus Ha­laple­mur) in Ra­nomafana Na­tional Park, Mada­gas­car. Intl J Pri­ma­tol, 20/4: 547-566.

Tan, C. 1999b. Life his­tory and in­fant rear­ing strate­gies of three Ha­pale­mur species. Pri­mate Re­port, 54/1: 33.

Wright, P., M. Ran­d­ri­manan­tena. 1989. Com­par­a­tive ecol­ogy of three sym­patric bam­boo lemurs in Mada­gas­car. Am J Phys An­thro­pol, 78: 327.

Ya­mashita, N., C. Tan, C. Vin­yard, C. Williams. 2010. Semi-quan­ti­ta­tive tests of cyanide in foods and exc­reta of three Ha­pale­mur species in Mada­gas­car. Amer­i­can Jour­nal of Pri­ma­tol­ogy, 72: 56-61.

Ya­mashita, N., C. Vine­yard, C. Tan. 2009. Food me­chan­i­cal prop­er­ties in three sym­patric species of Ha­pale­mur in Ra­nomafana Na­tional Park, Mada­gas­car. Am J Phys An­thro­pol, 139: 368-381.