MAMMALIAN SPECIES No. 148, pp. 1-7, 4 figs. LeOntOpitheCUS rOSalia. By Devra G. Kleiman Published 8 May 1981 by The American Society of Mammalogists Leontopithecus Lesson, 1840 Leontopithecus Lesson, 1840:184, 200. Type species designated by Pocock, 1917:255, as Leontopithecus marikina Lesson. Marikina Gray (not Lesson), 1843:xviii, 15. Type species by tau- tonomy Leontopithecus marikina Lesson. Leontocebus Elliot (part, not Wagner, 1840 = Saguinus Hoffmannsegg, 1807), 1913, l:xxxv, 194. Type species des- ignated as Hapale chrysomelas Wied (but Miller, 1912:380 had already designated Midas leoninas E. Geoffroy = Sa- guinus fuscicollis fuscus Lesson as type oi Leontocebus Wag- ner). Leontideus Cabrera, 1956:52. Renaming of Leontocebus Elliot. Type species Simia rosatia Linnaeus. CONTEXT AND CONTENT. Order Primates, Suborder Haplorhini, Infraorder Platyrrhini, Family Callitrichidae. In ad- dition to Leontopithecus, this family includes the extant genera Saguinus, Caltithrix, and Cebuella (Hershkovitz, 1972, 1977). The genus Leontopithecus includes only one living species (Hershkovitz, 1972; Coimbra-Filho and Mittermeier, 1972). Leontopithecus rosalia (Linnaeus, 1766) Lion tamarin [Simia] Rosatia Linnaeus, 1766:1, 41. Type locality Brazil, re- stricted to right bank, Rio Sao Jo?o, Rio de Janeiro, by Car- valho, 1965:22. Midas chrysomelas Kuhl, 1820:51. Type locality Ribeirao das Minhocas, left bank upper Rio dos llh?us. Southern Babia, Brazil. Jacchus chrysopygus Mikan, 1823:fasc. 3, pi. 1. Type locality Ipanema (=Varnhagen or Bacaetava), Sao Paulo, Brazil. [Jacchus] Chrysurus I. Geoffroy, 1827:520, attributed to Wied and cited as synonym of chrysomelas. [Jacchus Rosalia] Guyannensis Fischer, 1829:65. Type locality French Guiana (but Eastern Brazilian individual probably transhipped from Cayenne to France?Hershkovitz, 1977). [Jacchus Rosalia] Brasiliensis Fischer, 1829:65. Type locality not specified. [Midas] Leontopithecus marikina Lesson, 1840:200. New name for Simia rosalia Linnaeus. Midas [(Leontopithecus)] ater Lesson, 1840:203. Renaming of Jacchus chrysopygus Mikan. L[eontopithecus] aurora Elliot, 1913, 1:182. Lapsus for L. mari- kina Lesson. Leontocebus leoninus, Pocock (not Humboldt), 1914:898. CONTEXT AND CONTENT. Context in generic account above. The three kinds of Leontopithecus are currently consid- ered subspecies (Hershkovitz, 1972; Coimbra-Filho and Mitter- meier, 1973). The taxonomy of Leontopithecus has been dis- cussed by Hershkovitz (1977), Coimbra-Filho (1970a) and Coimbra-Filho and Mittermeier (1972). The following key to the subspecies is modified from Hershkovitz (1977): 1 Mane and trunk blackish, outer and inner side of thighs and rump contrastingly colored golden to reddish L. r. chrysopygus Mane, at least in front, mostly or entirely golden or red- dish, trunk blackish, reddish, golden or buffy, thighs not contrastingly colored 2 2 Trunk blackish L. r. chrysomelas Trunk reddish, orange, golden, or buffy L. r. rosalia The species includes three subspecies: L. r. chrysopygus Mikan, 1823, see above (ater Lesson a syn- onym). L. r. rosalia Linnaeus, 1766:1, 41, see above (guyannensis Fisch- er, brasiliensis Fischer, marikina Lesson, aurora Elliot, and leoninus Pocock are synonyms). L. r. chrysomelas Kuhl, 1820:51, see above (chrysurus Geoffroy a synonym). DIAGNOSIS. Lion tamarins are the largest of the caUi- trichids, with a long silky pelage. The face is almost bare and surrounded by a mane. The hands and digits are extremely long, with a web partially uniting the digits of the hand. GENERAL CHARACTERS. Weights of captive adult males and females average 710 g (range 600 to 800 g) in L. r. rosalia (Hoage, 1978, pers. comm.). There is no sexual dimor- phism, although females may average slightly heavier than males during pregnancy. Previously published weights are lower (Hershkovitz, 1977; Coimbra-Filho and Mittermeier, 1972; Napier and Napier, 1967), but some of these weights have combined juveniles and adults, as well as feral and captive-born animals (the latter are heavier). Length of head and body averages 261 mm (200 to 336) and length of tail averages 370 mm (315 to 400) (Hershkovitz, 1977; see also Coimbra-Filho, 1976; Coimbra-Filho and Mittermeier, 1972). The mane is derived from long hairs on the crown, cheeks, and throat and obscures the ears. Coloration of L. r. rosalia ranges from the pale golden to rich reddish-golden with occasional orange, brown, or black coloration on the tail and forefeet. L. r. chrysomelas and L. r. chrysopygus are predomi- nantly black. L. r. chrysomelas has golden to reddish-orange on the front of the mane, the lower half of the forelimbs and the proximal half of the dorsal surface of the tail. L. r. chrysopygus has varying amounts of reddish-golden coloration on the rump, inner and outer thighs, and base of tail. The glans penis is acorn- shaped (Hershkovitz, 1977). For more complete discussion of di- agnostic and general characters as well as color photographs of the three subspecies, see Hershkovitz (1977), Coimbra-Filho and Mittermeier (1973), and Coimbra-Filho (1976). A captive L. r. rosalia family at the National Zoological Park is shown in Fig. 1. DISTRIBUTION. Lion tamarins historically occurred in the lowland forests of southeastern Brazil, in the states of Babia, Espirito Santo, Rio de Janeiro, and Sao Paulo (Fig. 2). They are currently extinct in Espirito Santo (Hershkovitz, 1977; Coimbra- Filho and Mittermeier, 1973, 1977). L. r. chrysopygus was re- stricted to the tropical rain forests between the Rio Paranapa- nema and the Rio Tiet? in the state of Sao Paulo. Since most of the forest has been destroyed, the current range of this form is restricted to the Morro do Diabo State Park (in the westernmost part of Sao Paulo), with a second remnant population near G?lia, Sao Paulo (22?18'S, 49?34'W) (Hershkovitz, 1977; Coimbra-Filho, 1970a, 19706, 1976). L. r. chrysomelas has the most northerly range, and was originally found between the south bank of the Rio das Contas and the north bank of the Rio Belmente along the Atlantic Coast (14? to 16''S). Currently there are small, scattered populations near Buerarema, Itabuna, Una, and possibly llh?us, in the state of Babia (Hershkovitz, 1977; Coimbra-Filho and Mittermeier, 1973, 1977). Much of the original forest has been logged or has been replaced by cacao and rubber plantations, which are un- suitable for lion tamarins. L. r. rosalia was historically found in the coastal forests of the states of Rio de Janeiro (formerly Guanabara) and Espirito Santo (23?S, 44?W to 20.5?S, 40.5?W) south of the Rio Doce. The range is now reduced to remnant forests in the Rio Sao Joao Basin in Rio de Janeiro, which are scattered and probably total consid- erably less than 900 km^ in area (Coimbra-Filho and Mittermeier, 1977). It is not known how long the ranges of the three subspecies have been isolated. Both L. r. chrysomelas and L. r. rosalia live at altitudes of less than 300 m, while L. r. chrysopygus may range to 700 m. Distribution of L. rosalia is discussed further in Hersh- kovitz (1977), Coimbra-Filho (1970a, 19706, 1976), and Coimbra- Filho and Mittermeier (1973, 1977). There is no fossil record. MAMMALIAN SPECIES 148 FIGURE 1. A captive golden lion tamarin family at The National Zoological Park (Photo courtesy of Smithsonian Institution). FORM. Hershkovitz (1977) provided a complete summary of the morphology of L. rosalia, which forms the basis of this section unless otherwise specified. The remarkably different pat- tern of coloration of the three subspecies of L. rosalia has been attributed to a gradual evolutionary bleaching process (Hersh- kovitz, 1977). Individual differences in coloration and pattern in ^ L. r chrvsoDVQus S?O PAULO Rio Paranapanema I I 54? 51? 48' , Rio das Contas BAH?A \ > 300km\ ,, Rio /i^ "^ _ ?Una Rio Pardo ^^ j~ I6? ESPiRITO SANTO -20? RIO DE JANEIRO FIGURE 2. Past and present distribution of the three subspecies of Leontopithecus rosalia in Brazil. Based on Hershkovitz (1977) and Coimbra-Filho and Mittermeier (1977). Drawing by S. James. L. r. rosalia, and especially the tendency of captive golden lion tamarins to be nearly yellow-white in color, have often been at- tributed to diets which are deficient in carotenids and protein (Coimbra-Filho, 1965) or to excessive sunlight (Hershkovitz, 1977). However, captive L. r. rosalia acquire a rich, reddish- golden coloration from exposure to sunlight, which also causes a darkening of the facial skin (Kleiman, unpubl.). Coimbra-Filho and Cruz Rocha (1978) described a case of alopecia and depig- mentation in a specimen of ?. r. chrysomelas. Coimbra-Filho and Maia (1979a) described the molting patterns of captive L. r. ro- salia. The face of L. rosalia is naked, with variation in the degree of pigmentation; facial coloration ranges from fleshy to purplish. Hair on the forehead is short and wedge-shaped, unlike the mane which arises from long hair on the crown, cheeks, and throat. The external ear of L. rosalia is rounded in form, and becomes concealed as the mane develops in the juvenile. There is a median laryngeal air sac present which is larger in males than females (Hershkovitz, 1977). A sternal apocrine scent gland is present (Epple and Lorenz, 1967). The forelimbs are slender and elongate, and the long digits are designed for probing for grubs and insects. The fore and hindlimbs are more equal in length relative to other callitrichids. Limb proportions are as follows: length of humerus X 100/length of radius, 99 (98 to 103); length of femur x 100/length of tibia, 96 (94 to 101), length of humerus and radius X 100/length of femur and tibia, 88 (87 to 90). L. rosalia is less of a saltator than other callitrichids, with tendencies toward being a climber. The hand is shorter than the foot and the poUex is not opposable. The digits have sharp, recurved, laterally compressed claws. Relative lengths of digits, from longest to shortest, are: manus 3-4-2-5-1; pes 4-3-5-2-1. The palm is narrow, and the length of the middle finger is more than twice the width of the palm. The vertebral formula is C7, T12, L7, S3^, Cd22-28 (Hershkovitz, 1977). Two nipples are present and are axial in position. The milk of L. r. rosalia is similar to that of other New World monkeys (Buss, 1975). The penis is acorn-shaped and the penile stem is covered with papillae. The testes may be scrotal or inguinal, po- sition being under some voluntary control. Measurements of testes of two males were 10 X 9 and 15.5 X 11 m. The somewhat pendulous scrotum is unpigmented; greatest scrotal diameter for MAMMALIAN SPECIES 148 FIGURE 3. Photograph of skull of L. r. rosalia (courtesy of Smithsonian Institution). From top: dorsal and ventral views of cranium and ventral view of mandible. Scale is in mm. a male was 18 mm. The baculum of L. rosalia is the most spe- cialized of the callitrichids, with measurements of one individual being 3.0 mm long and 0.9 mm wide. Female external genitalia are unpigmented; the labia majora are more poorly differentiated than in other callitrichids. There are sebaceous and sudoriparous glands in the circumgenital area of both sexes (Hershkovitz, 1977). The skull of L. rosalia (Figs. 3, 4) has been described by DuBrul (1965) and Hershkovitz (1977). The dental formula is i 2/2, c 1/1, p 3/3, m 2/2, total 32. The upper and lower incisors are comparatively short, and the canines are twice as long as adjacent teeth. The relationship between incisor and canine lengths has resulted in the use of the term "long-tusked" to describe Leontopithecus and Saguinus and "short-tusked" for Callithrix and Cebuella, in which the canines and long incisors are approx- imately the same length. Upper incisors have expanded crowns, with the outer incisor set in an oblique manner and posterior to the inner incisor (Figs. 3, 4). The lower incisors also have trans- versely expanded crowns, with the lateral incisor club-shaped and larger than the spatulate-shaped central incisor. The upper and lower canines are long and saberlike. Permanent upper and lower premolars are more molariform than in other callitrichids. AU upper premolars are distinctly bicuspid with an incipient meta- cone often present in either or both P3 and P4. Lower premolars are also comparatively well molarized. Sample measurements (in FIGURE 4. Lateral view of cranium and mandible of skull shown in Fig. 3. mm) of an upper P2 and lower p2 respectively are: height of crown 3.0, 3.4; length 3.0, 2.9; width 3.3, 2.5. Ml is more than half again as large as the M2 and about a third larger than P4. The trigon basin of the Ml is deeply concave. Both upper molars have well-developed accessories. The lower molars are quadrate and simplified. The deciduous P4 is highly molarized which to- gether with the molarization of the permanent premolars supports the suggestion that the center of mastication has shifted forward (Hershkovitz, 1977). The greatest length of skull in adults is 50 to 62 mm. L. rosalia has four communicating sphenoidal cavitations opening behind in a pair of large sphenoidal fossae, which is unique among higher primates (Hershkovitz, 1977). The mandible outhne is in- termediate between the V-shape of Callithrix and the U-shape of Saguinus. The frontal contour is low and the nasal profile slightly concave. The orbits are somewhat square in outline with the interorbital area pneumatized. The auditory buUae are inflat- ed; the dental arcade is between a U- and V-shape. The brain of L. rosalia is relatively larger but less complicated than in other callitrichids. Four major sulci have been found on each hemi- sphere (Hershkovitz, 1977). Some aspects of the muscular system and internal organs of L. rosalia are given in Hill (1957). Be- nirschke and Richart (1963) provide weights of some internal or- gans. FUNCTION. Preliminary data on urinary chorionic gonad- otropin levels in pregnant females were presented in Kleiman et al. (1978). Bush et al. (1977) provided information on physiologic measures (blood pH and gases, rectal temperature, respiration, and pulse rates) during physical restraint of L. rosalia. ONTOGENY AND REPRODUCTION. As with other callitrichids, both fetuses share the same chorion (Benirschke and Layton, 1969). The placenta is discoidal with vascular anas- tomoses present, resulting in blood chimerism, a characteristic pe- culiar to callitrichids. Benirschke and Richart (1963) provided weights of fetuses and a placenta. Gestation was estimated as between 132 and 137 days, based on interbirth intervals (Hersh- kovitz, 1977; Rabb and Rowell, 1960; Ulmer, 1961); however, observed intervals between copulatory activity and birth suggest that gestation averages 129 days with a range of 125 to 132 days (Kleiman, 1977a; Wilson, 1977). Post-partum matings are most common approximately 3 to 10 days after birth (Snyder, 1974; MAMMALIAN SPECIES 148 Wilson, 1977). Detection of pregnancy is difficult without palpa- tion although the rise in urinary chorionic gonadotropins during weeks 4 to 9 of pregnancy suggests that the Non Human Primate Pregnancy test kit could be used to identify and chart the course of pregnancy (Kleiman et al., 1978). A behavioral estrus, with copulatory activity, is common during pregnancy shortly after the drop in chorionic gonadotropins (about 8 weeks pre-partum) (Klei- man and Mack, 1977). Otherwise, copulatory behavior is infre- quent and appears to be confined to the period near ovulation. The length of the ovarian cycle is unknown, and appears highly variable. However, 50% of observed intervals between peaks of sexual activity were from 14 to 21 days (Kleiman, 1977a), which is within the range of ovarian cycles reported for Callithrix jacchus by Hearn (1977). Litter size ranges from one to three, averaging two (Hershkovitz, 1977). In Brazil, births are concen- trated in a 6-month period between September and February, after the start of the rainy season (Coimbra-Filho and Mitter- meier, 1973; Coimbra-Filho and Maia, 19796). In the Northern Hemisphere births occur mainly from February to August (Klei- man, 1977a; Hershkovitz, 1977). Both captive and wild individ- uals in Brazil are rarely reported to bear more than one litter per year, while both wild-caught and captive-born females in the North Temperate Zone may have two litters per year (Kleiman, 19776). Neonates are fully furred and with the eyes opened. The fur is usually pale golden and there may be a black band extending frontoparietally which is later obscured as the mane develops. Birth weights average 60.6 g (range 52.1 to 74.6 g); singletons are heaviest and triplets hghtest (Hoage, 1978, pers. comm.). Neonatal length of head and body ranges from 100 to 114 mm and length of tail 117 to 143 mm (Hoage, 1978, pers. comm.). Growth rates may be found in Hoage (1978, pers. comm.), DuMond et al. (1979) and Rohrer (1979). Young cling to the mother from birth, and are able to crawl from her dorsal to ventral surface for nursing. During the first 2 weeks, infants are often asleep or nursing and only gradually increase their visual exploration and manipulation of the environ- ment. Manual and oral manipulation of objects increases rapidly from week 5. Infants first begin to leave carriers for brief periods during weeks 2 to 3 although well coordinated locomotion, such as jumping, leaping, and running is not common until week 8. Play activities become more common as locomotion and coordi- nation improve. Young begin to eat solid food from the parents' hands during week 4, and start to acquire food independently during week 8; weaning occurs by week 12. Young juveniles ac- quire considerable food from parents and older siblings through food sharing and stealing. From week 10 they begin occasionally to share food with relatives; the major increase in food sharing behavior by older juveniles, however, occurs at ages 25 to 35 weeks, when recently-born younger siblings begin to eat solid food. By week 16, young are approximately one-half of adult weight (Hoage, 1977, 1978; Green, 1979). Signs of sexual matu- rity, that is, sternal scent marking and back arching, appear be- tween weeks 32 to 40, and are more common in young males than females. Both sexes are sexually inhibited while still within the natal family group, and sibling or parent-offspring conflict may occasionally occur (Kleiman, 1979a). Most males and females are not sexually and socially mature until approximately 18 months of age (Kleiman and Jones, 1977). Other descriptions of devel- opment may be found in Ditmars (1933), Altmann-Sch?nberner, (1965), and Frantz (1963). The longevity of captive animals has increased with im- proved care; captive individuals between 10 and 15 years of age often are still reproductively active (Kleiman, 19796). ECOLOGY. The sparse information available on the ecol- ogy of L. rosalia may be found in Coimbra-Filho (1970a, 19706, 1976) and Coimbra-Filho and Mittermeier (1973, 1977). L. rosalia appears to be restricted to primary lowland forest, although little climax forest still exists within its range. Annual rainfall varies from 1100 to 2000 mm, with temperatures averaging 22?C. The wet season occurs from September to March within the range of L. r. rosalia and L. r. chrysopygus; in the range of L. r. chrys- omelas, heaviest rains fall from March to June and in November and December. L. rosalia is usually found between 3 to 10 m above the ground, in dense vines and epiphytes, which may pro- vide protection from aerial predators. Protection from pr?dation may also be improved by the tendency for groups to sleep in tree holes abandoned by other species. Entrances to these shelters have been found as low as 1.5 m from the ground although most are 11 to 15 m above the ground. Hair deposits found in these shelters suggest extended usage (Coimbra-Filho, 1977). Coimbra- Filho (1977) captured L. rosalia by trapping groups in tree holes. Lion tamarins feed mainly on fruits, insects and occasionally small vertebrates. Insects of the orders Blattaria, Orthoptera, Homoptera, Lepidoptera, Hemiptera, and Cole?ptera (especially larvae) are taken. Snails, lizards (Anolis spp.), and possibly bird eggs and hatchlings may also be eaten. Preferred fruits are soft, sweet and pulpy; species eaten include Ficus spp.. Inga spp., Tapirira guianensis, Cecropia sp., Posqueria latifolia, and sev- eral forms of Myrtaceae including Marlierea sp., Eugenia sp., and Campomanesia guabiroba. Group size varies from two to eight, with three to four being most common, although feeding aggregations of 15 to 16 have been reported. Groups are probably composed of a single repro- ductive pair, their offspring from one or two litters, and perhaps other relatives, thus forming a nuclear or extended family. There are no data on when juvenile dispersal occurs and the mecha- nisms of pair bond formation, although these phenomena have been examined in captivity (Kleiman 1978, 1979a). Home range size is not known. Lion tamarins are likely to be territorial, based- on the high levels of intrasexual aggression observed in neigh- boring groups in captivity (Snyder, 1974). L. r. chrysomelas may form mixed-species associations with Callithrix jacchus penicil- lata (Coimbra-Filho, 19706). The three subspecies of L. rosalia are near extinction because of human activity within their range, the major factor being the near total destruction of the primary forest. Current estimates of the population sizes of the three subspecies are: golden lion tam- arin (L. r. rosalia), about 100 to 200 animals; golden-headed lion tamarin (L. r. chrysomelas), about 200 animals; golden-rumped lion tamarin (L. r. chrysopygus), about 200 animals (Coimbra- Filho and Mittermeier, 1977). This species is in serious jeopardy, and may not survive. The export of specimens to zoos and as pets, especially L. r. rosalia, whose range is within one of the most densely populated areas of Brazil, may also have contributed to the species decline. Since the mid-1960's, conservation efforts have concentrated on three approaches: (I) the development of secure reserves, (2) the establishment of a captive breeding pro- gram in Brazil (the Tijuca Bank) to provide specimens for a r?in- troduction program, and (3) the implementation of an interna- tional cooperative breeding program to increase the captive population outside of Brazil. These topics are covered in several contributions in Bridgwater (1972) and in Hill (1970), Perry (1972), Magnanini (1977), Magnanini et al. (1975), Kleiman (19776), Klei- man and Jones (1977), and Coimbra-Filho and Mittermeier (1977). An International Studbook for L. r. rosalia, which docu- ments changes in the captive population, is prepared annually by D. G. Kleiman. Of the three conservation programs, the poorest success has been achieved in the establishment and protection of secure reserves within Brazil. Although reserves now exist for all three subspecies (Coimbra-Filho and Mittermeier, 1977), only the Morro do Diabo State Park may be sufficiently large to main- tain a viable population of L. r. chrysopygus without human in- terference. There are currently no firm estimates of population sizes within reserves for L. rosalia. However, Coimbra-Filho and Mittermeier (1978) do not recommend a reintroduction program in the near future. Captive breeding of the three subspecies at the Tijuca Biological Bank in Rio de Janeiro, directed by A. F. Coimbra-Filho, has been only partly successful, with many cap- tive births but also with losses from unknown causes (Coimbra- Filho, pers. comm.). Outside Brazil, breeding success has im- proved the status of the captive population of L. r. rosalia. Based on the International Studbook, the numbers of captive golden lion tamarins increased from 69 in 1972, to 74 in 1975, and to 120 in 1978 (excluding an additional 18 specimens in South Africa in 1978) (Kleiman, 19796) after a continuous annual decline in num- bers of captive specimens between 1968 and 1972, when Brazil imposed a ban on the species export (Perry et al., 1975). Care and maintenance techniques may be found in DuMond (1971, 1972) and Coimbra-Filho and Magnanini (1972). Hand-rear- ing and reintroduction techniques for hand-raised young may be found in DuMond et al. (1979) and Rohrer (1979). Captive spec- imens are susceptible to infection by Herpes viruses, rub?ola, the bacteria Pseudomonas, Klebsiella, Salmonella, and to infestation with parasites such as Prosthenorchis and Rictularia (Hershko- vitz, 1977). Due to its extreme rarity, L. rosalia has not been used in biom?dical research. BEHAVIOR. The following account is based mainly on observations of captive L. r. rosalia. L. rosalia is primarily ar- boreal. Individuals walk, run, and climb on branches using a quadrupedal gait. They frequently spring or leap from branch to MAMMALIAN SPECIES 148 branch and cling to and leap from vertical surfaces. In captivity, lion tamarins descend to the ground to explore and forage on the substrate. They apparently prefer to move in a horizontal plane (Coimbra-Filho and Magnanini, 1972). L. rosalia is diurnal, awak- ing near sunrise and retiring into nest boxes at dusk. A midday period of rest is common. The basic social group appears to be a nuclear or extended family, and intrasexual aggression among unfamiliar adults is high, especially if an animal of the opposite sex is also present (Ep- ple, 1967; Snyder, 1974). Females may be more aggressive than males (Snyder, 1974; Kleiman, 1979a). Within both artificially composed and natural family groups, only a single male and fe- male reproduce (Snyder, 1974; Kleiman, 1978), even though all members of a group may interact socially. Dominance interac- tions are infrequent within established groups, and outbreaks of aggression are unpredictable, often occurring when the repro- ductive female approaches estrus (Kleiman, 1979a). One sign of tension is the arch back display whose function and contextual occurrence have been analyzed by Rathbun (1979). Groups of lion tamarins exhibit frequent tactile contact with allogrooming and huddling being two major social activities. Allogrooming move- ments are typically primate in form, with the hands being used to part the fur and both hands and mouth used to remove detritus. Males groom females more than the reverse (Kleiman, 1977a). AUosniffing is commonly seen, with the male sniffing the female more frequently, especially in the anogenital region, as she ap- proaches estrus (Snyder, 1974; Kleiman, 1977a). Males also in- crease the frequency of other affiliative behaviors, such as groom- ing and huddling, as the female nears estrus. By contrast, femides exhibit an increase in affiliative behaviors directed towards males approximately 2 to 4 days before the males' peak sexual interest (Kleiman, 1977a). Copulatory behavior is primate in form with the male mount- ing and clasping the female in the mid-section anterior to the hindlimbs while keeping the hindfeet clasped on a branch or other surface. Shallow rapid thrusting movements precede intromis- sion. Several intromissions may precede an ejaculation. Ejacu- lation is difficult to discern but may be recognized by (a) the female vocalizing and/or looking back to the male, (b) a prolonged thrust during intromission, (c) licking of the genitalia by the male and female after separation, and (d) a quiescent period without attempted mounts following separation (Snyder, 1974; Kleiman, 1977a). Copulatory activity, although infrequent, is more common in newly established pairs than in pairs with long-term bonds and with offspring (Kleiman, 1977c). Both males and older juveniles participate in parental care activities (Hoage, 1977, 1978, pers. comm.; Frantz, 1963; Alt- mann-Sch?nberner, 1965; Snyder, 1974; Green, 1979). Immedi- ately post-partum, the female carries the neonates exclusively and may be aggressive to family members who approach (Hoage, 1977; Snyder, 1974). Between 2 and 17 days post-partum, the young are first transferred to the male, with triplets usually being transferred earlier than twins or singletons (Hoage, 1977; Klei- man, 1977c). First transfers are accomplished by persistent in- terest by family members who approach, press the ventrum against the young, and manipulate young with the hands. Such transfers are facilitated by the mother's developing intolerance, expressed by scratching at the infants, and rubbing her back (young are carried dorsally) against surfaces. Hoage (1977, 1978) reports that: (1) mothers are the principal carriers of young through week 3, with fathers carrying more than mothers there- after; (2) juvenile females begin carrying young earlier (week 2) than juvenile males (week 3), but juveniles of both sexes rarely carry younger siblings after week 8; (3) there is a tendency for parents (and possibly older juveniles) to preferentially carry in- fants of their own sex; and (4) primiparous parents with previous exposure to infants are most successful in rearing their own neo- nates, suggesting that experience with infants during the juvenile phase is important for successful reproduction. Hoage (1978; pers. comm.) charted the social development of young lion tamarins, especially the ontogeny of food-sharing, an unusual behavior first reported by Wilson (1976) and further analyzed by Brown and Mack (1978) and Green (1979) with respect to the accompanying visual and vocal signals. Play consists main- ly of chasing, grappling, and wrestling, and is observed primarily in juveniles (Snyder, 1974; Hoage, 1978, pers. comm.). Visual signals are not as complex as in Old World monkeys, but include agonistic patterns such as arched back displays (Rathbun, 1979), stares, open mouth threats, and piloerection. Tongue-flicking is a display seen mainly in the context of sexual behavior (Epple, 1967; Snyder, 1974). Vocalizations are complex; they con- sist of whines, trills, clucks, and non-tonal sounds, which can be combined to form more complex calls (McLanahan and Green, 1977; Green, 1979). There are few sex differences in the frequency or types of vocalizations. Of special interest is the Long Call, composed of trills, whines, and clucks, which bonded pairs produce as a duet. This call may function in spacing groups in the wild. Green (1979) and McLanahan and Green (1977) ana- lyzed the contextual occurrence of the different vocalizations, and Green (1979) described vocal ontogeny. Olfactory communication is mainly accomplished through scent marking with the sternal and circumgenital glands (Epple and Lorenz, 1967; Epple, 1972). Adult reproductive males and females scent mark at similar frequencies; young females rarely scent mark until paired, but young males begin scent marking while still in the natal family group. Adult reproductive males scent mark at relatively constant frequencies; breeding females, on the other hand, exhibit fluctuations, including low levels at estrus and after birth and higher levels during pregnancy (Mack and Kleiman, 1978; Kleiman and Mack, 1980). GENETICS. The diploid number of chromosomes is 46. There are 4 metacentric, 28 submetacentric, and 12 acrocentric chromosomes. The X chromosome is submetacentric and the Y is metacentric (Hsu and Hampton, 1970). Five hybrids of L. r. rosalia and L. r. chrysomelas from three litters (none of which survived) were intermediate in color pattern between the parental types, although the color pattern of each hybrid differed (Coim- bra-Filho and Mittermeier, 1976). Captive-born L. rosalia have been found with defects of the diaphragm, resulting in the mi- gration of liver, stomach, and intestines into the chest region. The distribution of the defect in the captive population strongly suggests a genetic basis, but it is not the result of inbreeding of captive individuals (Bush et al., in press). REMARKS. L. r. rosalia is often referred to as the golden marmoset. Other common names for the species are maned mar- moset and silky marmoset. I am grateful to C. Handley and R. Thorington for assistance with the synonymies. S. James pre- pared the distribution map, and V. Garber and G. Hill typed the manuscript. LITERATURE CITED Altmann-Sch?nberner, D. 1965. 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