Saguinus mystax (Spix, 1823)
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https://doi.org/ 10.5281/zenodo.5730714 |
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https://doi.org/10.5281/zenodo.5730874 |
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persistent identifier |
https://treatment.plazi.org/id/DF668780-FFC9-FFD8-FFC7-FEE8696EE64F |
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Conny |
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Saguinus mystax |
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Mustached Tamarin
French: Tamarin 8 moustaches / German: Schnurrbarttamarin / Spanish: Tamarin bigotudo Other common names: Moustached Tamarin; Red-capped/Red-cap Mustached Tamarin (pileatus), Spix's Mustached Tamarin (mystax), White-rump Mustached Tamarin (pluto)
Taxonomy. Midas mystax Spix, 1823 View in CoL ,
between Rio Solimoes and Rio Ica, Amazonas, Brazil. Restricted by P. Hershkovitz in 1977 to Sao Paulo de Olivenca .
C. P. Groves in his book Primate Taxonomy published in 2001 found that, while the subspecies mystax and pluto are very similar to each other, the form pileatus is quite distinct, and he listed it as a separate species. Nevertheless, current evidence indicates that distributions of mystax and pluto are separated by pileatus, and as such, pileatus is not recognized as a separate species here. Reports in the 1980s of S. mystax occurring in northern Bolivia have not been confirmed. Three subspecies recognized.
Subspecies and Distribution. S. m. mystax Spix, 1823 — S of the Amazon River in WBrazil (W of the Rio Jurua) and NE Peru, as far W as the E banks of the middle and lower Rio Tapiche and the lower Rio Ucayali, further S it extends W as far as the Rio Ucayali, crossing the Rio Tapiche at c.6° 40’ S, the S limit is marked by the rios Urubamba and Inuya. S. m. pileatus 1. Geoffroy Saint-Hilaire & Deville, 1848 — Brazilian Amazon, W of the Rio Purus, S at least as far as the Rio Pauini or Rio Mamornia. S. m. pluto Lonnberg, 1926 — Brazilian Amazon, from the W (left) bank of the Rio Purus, N of the Rio Tapaua, as far W as the Rio Coart. View Figure
Descriptive notes. Head—body 24-2 cm (males, n = 75) and 24-6 cm (females, n = 60), tail 38-2 cm (males, n = 72) and 38-7 cm (females, n = 57); weight 380-630 g (males from three sites, n = 161) and 360-650 g (females from three sites, n = 104) (single measurements are means from the nominate subspecies mystax , “Spix’s Mustached Tamarin”). Female Mustached Tamarins are a little larger than males. At the Rio Blanco, Peru, mean weight of 16 adult males was 564-5 g, with a range of 502-681 g. In Spix’s Mustached Tamarin, the skin around the mouth and nostrils is unpigmented pink and covered with white hairs, giving the characteristic mustached appearance. The mustache is distinctly larger than that of the Red-bellied Tamarin (S. labiatus ). Hairs on the middle of the chin are black. All three subspecies of Mustached Tamarin have long hairs on the mantle, shoulders, and upper arms that are brown or blackish, usually with a very fine silvery or buffy ticking, but not defined from the middle back. In Spix’s Mustached Tamarin, the crown and tail are black. The mantle is blackishbrown lightly ticked with orange. Arms, legs, hands, and feet are black. Hairs of the back, rump, and outersides of the thighs are blackish with orange ticking. The throat, chest, belly, and inner sides of limbs are blackish brown. The tail is black except at the base, which is ticked with orange like the rump. The tip of the tail is sometimes gray. External genitalia are unpigmented and covered with white hairs. Spix’s Mustached Tamarin can be distinguished from the other two subspecies by the dominantly blackish or dark brown color of the head, trunk, and extremities. The “Red-capped Mustached Tamarin” (S. m. pileatus) is distinguished from the other subspecies byits rustyred crown that extends forward onto the forehead in a thin line between the eyes. The rest of the face is black except for the typical white moustache. The skin around the mouth and nostrils is pink and unpigmented. The mantle is blackish brown, with fine buffy or orange ticking, with the paler base of the hairs showing through. The middle and lower back and rump have mixed blackish brown and buffy hairs. Outer sides of arms and legs are brown, becoming blacker on the hands and feet. The entire ventral surface and innersides of limbs are blackish brown. The tail is black. External genitalia are mostly or entirely unpigmented and covered with white hairs. The “White-rump Mustached Tamarin” (S. m. pluto) is similar to Spix’s Mustached Tamarin, except for the depigmentation of the anogenital and inguinal region and base of the ventral surface of the tail, which are covered with white hairs, hence its common name. The rest of the tail and the crown of the head are black. The area around the mouth and nostrils is unpigmented and covered in white hairs and whiskers, forming the moustache and extending out from the cheeks but with a small black patch on the middle of the chin. The mantle is black and ticked with buff (bases of the hairs are drab, sometimes whitish). Arms and upper surfaces of hands and feet are black. The lower back and outer sides of thighs are like the mantle but with more buff showing through. The throat, chest, and underparts are blackish brown.
Habitat. Tall and secondary lowland rainforest, preferring dense vegetation in secondary forest, tree falls, and edge habitat. Mustached Tamarins only marginally occupy seasonally flooded and inundated forests, possibly because they are more dependent on a continuous supply of fruit than sympatric species of saddle-back tamarins. Fruiting is more seasonal in the inundated forests than on the terra firma or alluvial plain forests, which often lack fruit during the dry season. The saddle-back tamarins are smaller and can resort more easily to gums and nectar during times of fruit scarcity. The Mustached Tamarin rarely forages along tree trunks (a typical foraging site for saddle-backs) and spends less time foraging in the understory where the saddle-backs are most active. Mustached Tamarins prefer the upper layers of the forest canopy. Spix’s Mustached Tamarin (subspecies mystax ) is sympatric with Geoffroy’s Saddleback Tamarin (S. nigrifrons ) and Spix’s Saddle-back Tamarin (S. fuscicollis fuscicollis ). The Red-capped Mustached Tamarin (subspecies pileatus) and the White-rump Mustached Tamarin (subspecies pluto) are sympatric with Spix’s Saddle-back Tamarin (S. J. avilapiresi) throughout their known distributions. The natural history of mustached tamarin is now quite well known. Spix’s Mustached Tamarin has been studied at sites in the Peruvian Amazon: on the left bank of the Rio Blanco, a tributary of the Rio Tahuayo (in turn a tributary of the Rio Amazonas, a little upstream from Iquitos) at Estacion Biologica Quebrada Blanco 1 (M. Ramirez, M. Norconk, and P. Garber) and on the right bank of the Rio Blanco at Estacion Biologica Quebrada Blanco 2 (R. Castro, T. Fang, E. Heymann, P. Lottker, M. Huck, A. Smith, and others). P. Soini and M. de Soini studied demographics of wild populations at sites in forests along the rios Maniti, Tahuayo, and Tapiche in north-eastern Peru. Behavioral, ecological, and demographic studies have also been done on Padre Isla, a seasonally flooded island in the Rio Amazonas near Iquitos, Peru (L. Moya, C. Gazzo, P. Garber, and E. Heymann ). Formerly devoid of primates , Spix’s Mustached Tamarin was introduced there by the Proyecto Peruano de Primatologia from 1977 to 1980: eleven individuals in 1977, 17 individuals in 1978, and 65 individuals in 1980. Mixed-species groups of Red-capped Mustached Tamarins and Spix’s Saddle-back Tamarins were studied at a site along the Rio Urucu by C. Peres.
Food and Feeding. Spix’s Mustached Tamarin at Quebrada Blanco feed on a wide range of fruits (c.90 species), plant exudates (nectar and gum), and invertebrates, including insects, especially Orthoptera , along with small frogs and lizards. During a seven-month period (June-December; late wet season and early wet season punctuated by a month of dry season in August) at the Rio Blanco, the diet of a group of Spix’s Mustached Tamarin was 42-3% fruits, 40-4% insects, 9:3% legumes, 5-6% nectar, and 2:2% gums. The percentage of feeding time devoted to fruits ranged from 37-6% in the dry season to 48:6% in the wet season. Nectar is an important component of the diet in the dry season when fruits are scarce. During dry season months (July=August) at Quebrada Blanco, mixed groups of Spix’s Mustached Tamarin and Geoffroy’s Saddle-back Tamarin devote 22-31% of their feeding time to nectar from the red flowers of Symphonia globulifera (Guttiferae) trees. A 14-month study of gum feeding at Quebrada Blanco showed that Spix’s Mustached Tamarins exploited fewer species (seven) than Geoffroy’s Saddle-back Tamarins (17 species), but Parkia nitida and P. igneiflora ( Fabaceae ) were extremely important to both during the dry season. During a full year, the diet of a group of Mustached Tamarins introduced to Padre Isla was 43-7% fruit, 34-9% animal prey, and 19-6% gums. The main families providing fruits were Fabaceae (especially Inga ) and also Moraceae . Stryphnodendron (Fabaceae) was an important source of gum. At the Urucu site, a mixed-species group of Redcapped Mustached Tamarin and Spix’s Saddle-back Tamarin ate ripe fruits form 155 species of plants. During 13 months, the diet of the Red-capped Mustached Tamarin was 70-6% fruits, 12:4% animal prey, 10-4% gums (mostly Parkia nitida and P. pendula), 6-2% nectar, and 0-1% unripe seeds and mushrooms. Although fruits dominate the plant part of diet most of the year at Urucu, nectar (especially of Symphonia globulifera) and gums were important during the late wet and early dry seasons in July-September. During the year, the most important genera and species providing fruits included Ficus (FE trigona, F. mathewsu, F. guianensis), Pseudolmedia laevis ( Moraceae ), Inga (Fabaceae) , Souroubea guianensis ( Marcgraviaceae ), Pourouma formicrum ( Urticaceae ), Hebepetalum humiriafolium ( Linaceae ), Leonia glycycarpa ( Violaceae ), and Minquartia guianensis ( Olacaceae ). Mustached Tamarins disperse seeds from a large number of lianas and trees. Some of the seeds they swallow are large, up to 2-5 cm in length. They are rapidly passed through the digestive tract (one to five hours, but usually less than three hours) and many of them are excreted more than 200 m from the parent tree. Heavier seeds are excreted more rapidly than lighter seeds. Mustached Tamarins have also been seen, on very rare occasions, eating soil (geophagy). The reasons for this are not known, but it may be an important mineral supplement, perhaps for certain trace elements. Large orthopterans (katydids, Tettigoniidae ) are preferred prey, but they also eatstick insects (Phasmatodea), praying mantises ( Mantodea ), lepidopteran eggs, cuckoo-spit aphids ( Hemiptera ), weevils, spiders, spider eggs, and scorpions. Foraging techniques of Mustached Tamarins and places they search are notably distinct from the more specific sites (crevices and knotholes) that saddle-back tamarins forage in. Mustached Tamarins travel and forage in the middle and lower canopy above the saddle-back tamarins. Both Mustached Tamarins and saddle-back tamarins forage in (and drink water from) bromeliads. Overlap in prey species captured by Mustached Tamarin and Geoffroy’s Saddle-back Tamarin can be minimal. In one study, only three of the 35 prey species identified were eaten by both (9:4% overlap). In a second study, however, overlap was higher; 31 of 62 prey species that could be identified were captured only by Geoffroy’s Saddle-back Tamarin, ten only by Spix’s Mustached Tamarin, and 21 eaten by both (34% overlap). Differences in prey selection reflect different foraging techniques. Mustached Tamarins, foraging in the foliage, find and eat generally smaller insects, most of them green, than those taken by saddle-back tamarins. Vertebrate prey includes mostly frogs ( Osteocephalus and Phyllomedusa , Hylidae ) but also some lizards (e.g. Norops, Polychrotidae ).
Breeding. Breeding is usually restricted to a single female, but groups generally contain more than one adult male; the male-female ratio in eight groups studied in the wild was 1-4:1. In four of the groups, paternity was monopolized by one male, but different males sired offspring in successive years in another group. In another group, the twins were believed to have been sired by two males. Relatedness in groupsis generally high, and most non-breeding individuals are either born in the group they are in or are closely related to the same sex breeder. It is unusual for groups to have individuals unrelated to any of the other group members. Births occur throughout the year, except for the dry season months ofJune and July. There is a birth peak during the wet season in November—February, when fruit and probably insect availabilities are at their highest. With a gestation of 150-160 days, the mating season is accordingly in July-September. The norm is for just one litter of twins per year. Interbirth intervals are 8-14 months and average 11-12 months. One study estimated 54 and 64-82 days of postpartum ovarian inactivity in two females, which is 3—4 times longer than in other callitrichids. Mate guarding by males has been observed when females are believed to be ovulating. The stimulus for this mate-guarding is probably from the female’s scent marks. Unlike female saddle-back tamarins, female Mustached Tamarins show physical signs of receptivity with conspicuous cyclic swelling of the vulva (turgid, moistened, and reddish), swollen nipples, and an increase in the size and activity of the suprapubic gland.
Activity patterns. The daily activity budget of a group of Red-capped Mustached Tamarin at Urucu was 30% moving, 26% feeding, 20-6% foraging for animal prey, 16-6% resting, and 6-8% engaged in social behavior, intergroup encounters, and other activities. The group’s active day averaged nine hours and twelve minutes. Although they spend much of the day traveling together, Mustached Tamarins and saddle-back tamarins do not sleep together. They separate in the late afternoon and sleep in different trees. They spend over 80% of their time within 50 m of each other and 72% of their time within 20 m of each other. Respective groups at Urucu slept in sites that were 15-120 m apart. When going to their sleeping sites, usually c.16:00 h, they become very quiet, travel rapidly, and follow each other, often single file, along the same pathways. Mustached Tamarins use five types of sleeping sites: the palm Jessenia bataua, tree hollows, dense tangles of foliage in the crowns of trees or near the trunks, crotches of large branches, and horizontal branches in tree crowns. They mostly use palm trees or dense tangles of foliage and occasionally tree crotches and open branches. In contrast, saddle-back tamarins, while preferring palms, use tree hollows as a second choice, but never sleep in crotches or along horizontal branches in the open. Mustached Tamarins do not use tree hollows. Although both species sleep in the same general locations on different nights, neither use specific sleeping sites used by the other species. Mustached Tamarins sleep higher up, for example, in taller palms. Their sleeping sites tend to be in the core of their home range, with no overlap with other conspecific groups.
Movements, Home range and Social organization. Spix’s Mustached Tamarins are generally found in groups of ¢.5-6 individuals, but groups of 9-10 individuals have been observed. Number of adults in each group is 2-6, usually with more adult males than females. The group studied at Rio Urucu had 8-10 individuals over a year, with up to 3-5 adult males and 1-2 adult females. When a group has more than one adult female, they are usually of different ages. Groups are quite stable, with reproductive pairs maintaining tenure for four to five years and even as many as seven. Migrations tend to occur following stability-disrupting events, such as entry of new individual into the group when one of the breeding pair is lost, or when groups become too large. Breeding and non-breeding males and females may emigrate to other groups or form new groups. Home ranges of groups at the Rio Blanco were 40-45 ha, with daily movements of 1500-1946 m. The home range of a mixed-species group of Redcapped Mustached Tamarin and Spix’s Saddle-back Tamarins at the Rio Urucu was much larger, ¢.179 ha, but with similar daily movements of 1150-2700 m/day. Aggressive interactions between group members are rare, but they tend to occur over food sources and during intergroup encounters when adult males behave aggressively toward the adult female, preventing her from approaching members of the neighboring group. Grooming is an important feature of their daily social interactions, and it reflects different social standing of group members; the adult reproductive female, for example, is consistently groomed more than she grooms others. Associated groups of Mustached Tamarins and saddle-back tamarins defend their home ranges jointly, with encounters between groups involving vigilance, vocal battles, chases, and even physical combat. Males and females participate, and aggression is directed principally at individuals of the same sex and species. Encounters can last up to two hours. Competition for food resources within the mixed-species groups is not critical, largely due to differences in theirsize, selection of animal prey, and dietary emphasis; saddle-backs, for example, exploiting more small trees and shrubs in the understory. Competition for key fruiting trees among groups of the same species, however, is important. The ability of groups to defend their home ranges is dependent on group size, and this might be a reason for the association between the different species. Predation of a female Spix’s Mustached Tamarin by an anaconda ( Eunectes murinus) was observed at Padre Isla; the anaconda caught her when her group was crossing a narrow lake using fallen tree trunks as stepping-stones. Mustached Tamarins mob predators and may even attack them. A mixed-species group of Geoffroy’s Saddle-back Tamarins and Mustached Tamarins were seen mobbing a 1-5-m boa constrictor ( Boa constrictor) when it caught a subadult male Mustached Tamarin by his leg. An adult male Mustached Tamarin bit the boa constrictor just behind its head and a female did the same about halfway along its body length. Their attacks were successful and caused the boa constrictor to release the subadult and leave the area. Crested eagles (Morphnus guianensis) are also predators of Mustached Tamarins. Alarm calls for avian predators are given at a rate of ¢.0-3 calls/hour—the majority elicited by raptors, and the rate of alarm calling increases following the birth of an infant. On hearing or giving an alarm call, Mustached Tamarins climb down or run from the periphery of a tree to the trunk, occasionally hiding under large leaves of palms. Other birds that cause alarm include ornate hawkeagle (Spizaetus ornatus), vultures (Coragyps, Cathartes), forest falcons (Micrastur), blackcollared hawks (Busarellus nigricollis , an insect eater that often follows tamarins), herons, yellow-rumped caciques (Cacicus cela), and smooth-billed anis (Crotophaga ani). Toucans (Ramphastos) and species of parrots can also elicit alarm calling. Densities of Spix’s Mustached Tamarin at 18 localities in Peru ranged from 7 ind/km? (Rio Tamshiyacu) to 84 ind/km? (Quebrada Vainilla, Rio Amazonas), or 14-14 groups/km?. Most densities were 20-30 ind/km? or 5 groups/km?. Surveys at three localities in the Rio Jurua Basin in Brazil provided estimates of 22 ind/km?, 31 ind/km?, and 33 ind/ km?. Densities of Red-capped Mustached Tamarins at threesites in and near the Urucu Basin in Brazil were 10-7 ind/km?, 11-6 ind/km?, and 12 ind/km?.
Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Mustached Tamarin is generally common but less adaptable to environmental disturbance than other tamarins. It is sometimes hunted for meat by local people because of it large size. There are two protected areas within the distribution of Spix’s Mustached Tamarin in Brazil: Serra do Divisor National Park andJutai-Solimoes Ecological Reserve. The only protected area in the Peruvian part of its distribution is ACR Comunal Tamshiyacu-Tahuayo. The White-rump Mustached Tamarin may occur in the part of the Abufari Biological Reserve thatis west of the Rio Purus.
Bibliography. Aquino & Encarnacion (1994b), Box & Morris (1980), Fang (1990), Garber (1986, 1988a, 1988b, 1993a, 1993b), Garber & Dolins (1996), Garber & Kitron (1997), Garber & Pruetz (1995), Garber & Teaford (1986), Garber, Encarnacion et al. (1993), Garber, Moya & Malaga (1984), Garber, Moya, Pruetz & Ique (1996) , Garber, Pruetz & Isaacson (1993), Gazzo (1990), Glander et al. (1984), Gozalo & Montoya (1992), Hershkovitz (1977), Heymann (1987a, 1987b, 1990a, 1990b, 1990c, 1992a, 1992b, 1993, 1995a, 1995b, 1996, 1997, 1998b, 2000b, 2000c, 2001), Heymann & Buchanan-Smith (2000), Heymann & Hartmann (1991), Heymann & Sicchar Valdez (1988), Heymann & Smith (1999), Heymann et al. (2000), Huck, Lottker & Heymann (2004a, 2004b), Huck, Lottker, Heymann & Heistermann (2004), Knogge & Heymann (2003), Lottker (2005), Lottker, Huck & Heymann (2004), Lottker, Huck, Heymann & Heistermann (2004), Moya, Ique & Soini (1990) , Moya, Verdi et al. (1990) , Nickle & Heymann (1996), Norconk (1990), Oversluijs & Heymann (2001), Peres (19913, 1992a, 1992b, 1993a, 1993b, 1993c, 1993d, 1994a, 1994b, 1996b, 2000b), Ramirez (1984, 1985, 1989), Rylands & Mittermeier (2008), Rylands et al. (1993), Sicchar & Heymann (1992), Smith (1997, 2000a, 2000b), Smith, Knogge et al. (2007), Smith, Tirado et al. (2001), Snowdon & Hodun (1985), Snowdon & Soini (1988), Soini & de Soini (1983, 1986, 1990a, 1990b), Soini et al. (1989), Tapia et al. (1990), Tello et al. (2002), Tirado et al. (2003).
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