Callicebus cupreus (Spix, 1823)

Voss, Robert S. & Fleck, David W., 2011, Mammalian Diversity And Matses Ethnomammalogy In Amazonian Peru Part 1: Primates, Bulletin of the American Museum of Natural History 2011 (351), pp. 1-81 : 43-46

Callicebus cupreus (Spix, 1823)

VOUCHER MATERIAL (TOTAL 5 16): Nuevo San Juan (AMNH 268239; MUSM 11116, 11117), Orosa (AMNH 73703–73708), Quebrada Esperanza (FMNH 88858–88860), San Fernando (FMNH 88856, 88857), Santa Cecilia (FMNH 86989, 86990).

UNVOUCHERED OBSERVATIONS: Actiamë ( Amanzo, 2006), Choncó ( Amanzo, 2006), Itia Tëbu ( Amanzo, 2006), Jenaro Herrera ( Aquino, 1978), Reserva Comunal Tamshiyacu-Tahuayo ( Puertas and Bodmer, 1993; Heymann and Aquino, 1994), Río Orosa ( Freese et al., 1982), Río Tapiche ( Bennett et al., 2001), Río Yavarí (left bank below Angamos; Salovaara et al., 2003), Río Yavarí-Mirím ( Salovaara et al., 2003), Tapiche ( Jorge and Velazco, 2006). (Note that some reports of Callicebus from the Yavarí-Ucayali interfluve have identified the local species as C. caligatus or C. moloch ; see below.)

IDENTIFICATION: The Amazonian forms of Callicebus (titi monkeys) were revised by Hershkovitz (1963), who recognized only two valid species, C. moloch and C. torquatus . However, Hershkovitz (1990) and Groves (2001) subsequently recognized 12 Amazonian species, and Roosmalen et al. (2002) recognized 22 Amazonian species that they assigned to four species groups. 13 Although most taxonomic diagnoses in this literature are based on coat-color differences, there is some evidence that the species groups recognized by Roosmalen et al. (2002) are morphometrically differentiated and have distinctive karyotypes ( Kobayashi, 1995). Unfortunately, the karyotypes of many alleged species are unknown, and molecular evidence

supporting the recognition of distinct taxa within species groups is almost nonexistent.

Hershkovitz (1963) referred all of the Callicebus specimens he examined from the Yavarí-Ucayali interfluve to the subspecies C. moloch cupreus , but he later ( Hershkovitz, 1990) identified some specimens of titi monkeys from the Yavarí-Ucayali interfluve as C. cupreus and others from the same region as C. caligatus . A small series of AMNH specimens from Orosa was listed among the material examined for both species in the latter publication, and an accompanying map ( Hershkovitz, 1990: fig. 36) showed C. cupreus and C. caligatus as occurring sympatrically at Orosa. Later, Hershkovitz (personal commun., 1995) explained that this was an error caused by listing original and revised identifications of the same specimens in separate species accounts.

In fact, titi specimens from the Yavarí- Ucayali interfluve could be referred either to Callicebus cupreus or to C. caligatus as those species were diagnosed by Hershkovitz (1990). For example, the forehead is grayish or reddish agouti with only a narrow blackish brow-line (as described for cupreus ) in most of our specimens (e.g., AMNH 73708, 268239), but in several others (e.g., AMNH 73705, 73706) the entire forehead and lower crown is blackish (as described for caligatus ). Similarly, some of our specimens (e.g., AMNH 268239) have clear reddish hands, feet, and lower limbs (as in cupreus ), whereas others (e.g., AMNH 73703, 73706) have dark reddish-brown or even blackish extremities (as in caligatus ).

According to Roosmalen et al. (2002), only Callicebus cupreus inhabits the Yavarí- Ucayali interfluve, but some of our specimens more closely resemble their diagnosis of C. caligatus , particularly with respect to the presence of a blackish forehead and lower crown as described above. Additionally, the basal part of the tail is blackish in some of our specimens (e.g., AMNH 73703) and others (e.g., AMNH 268239) have extensively pale-tipped tails, both traits associated with C. caligatus rather than C. cupreus in their diagnoses. In effect, our material combines traits of both species as recognized by these authors, who do not list any material examined, and whose illustrations do not correspond closely with their text. 14

We are not convinced that Callicebus caligatus or any of the other nominal taxa belonging to the Cupreus Group of Kobayashi (1995) and Roosmalen et al. (2002) — including C. caquetensis , C. discolor , C. dubius , C. ornatus , C. stephennashi , and probably C. aureipalati — are valid species distinct from C. cupreus . Although these forms collectively range across most of western Amazonia, they resemble one anoth- er in pelage markings (see Roosmalen et al., 2002: fig. 49; Defler et al., 2010: fig. 5; Wallace et al., 2006: figs. 4, 5), and at least some allegedly diagnostic differences seem to break down when large series of specimens (as opposed to just a few exemplars) are examined. Additionally, nonpelage differences among these taxa appear to be wanting: all analyzed forms are morphometrically similar ( Kobayashi, 1995), have the same diploid number of chromosomes (2 n 5 46; Roosmalen et al., 2002; Defler et al., 2010), and seem to occupy the same ecological niche (inhabiting forest edges and disturbed vegetation; Roosmalen et al., 2002). Because no verifiable examples of sympatry are known among nominal species in this group, it is plausible that these taxa represent nothing more than geographic coat-color variants of the same biological species, a hypothesis that merits future evaluation with molecular sequence data and with targeted collecting in areas where different coat-color phenotypes might be expected to come in contact.

External and craniodental measurements of Callicebus cupreus specimens that we examined from the Yavarí-Ucayali interfluve are summarized in table 14 . Most specimens are unaccompanied by weight data, but an adult male (MUSM 11116) from Nuevo San Juan weighed 1020 g, and an adult female (AMNH 268239) from the same locality weighed 1042 g.

ETHNOBIOLOGY: The Matses call the titi monkey wade¨, which is not linguistically

analyzable. It has three other supposedly ancient synonyms, tsokon, sëde¨, and masoko. The first synonym is onomatopoetic and unique to Matses. The other two are of uncertain etymology: both occur in neighboring Mayoruna languages, and therefore could easily be either borrowings or truly archaic words. Some Matses assert that masoko is not archaic, but a borrowing; there is no controversy about the other two names. Two subtypes are recognized, wadë piu ‘‘red titi monkey’’ and wadë çhëşhë ‘‘black (dark-colored) titi monkey.’’ The first is the commonly seen variety, which is found close to rivers. The second variety is reported to live in upland forest far from rivers, where the red variety is absent. Not all Matses hunters are familiar with the darker variety or accept that two varies exist. Those that claim to have seen the darker variety do not know much about it. (Despite a considerable effort to find it, the darker variety was not collected during our mammal inventory at Nuevo San Juan; it probably represents another case of overdifferentiation of a single biological species.)

Matses eat titi monkeys, but because they are small, hunters usually do not make a special effort to find them. Traditionally, as larger game became scarce in a locality, more titi monkeys would be killed. Nowadays, most hunters will not expend a shotgun shell to kill one despite the shortage of game around the larger and more permanent villages. One way to kill titi monkeys is to simply shoot them from the ground. Another way is to climb up at tree where the titi monkeys are hiding. When the titi monkeys see the hunter up in the tree, they will usually drop and flee running on the ground, where waiting hunting dogs and/or a second hunter will chase them down and kill them. Matses do not imitate their call to hunt them.

As with all monkeys, titi monkey are dressed for the pot by singing off the fur and then gutting them. Unlike large monkeys, titi monkeys (and other small monkeys) may be roasted instead of boiled. Pregnant women or men with pregnant wives cannot eat titi monkeys, lest their children grow up to have bad teeth [titi monkeys have black teeth]. Young people will not eat the liver of a titi monkey, lest it make their teeth rot.

MATSES NATURAL HISTORY: Titi monkeys have red underparts, a black brow, black teeth, and a whitish tail. They are small, the size of a squirrel monkey.

Titi monkeys can be found in any forest type, but they strongly prefer habitats that have open canopies [i.e., where light shines through to the understory], such as abandoned swiddens, riverside forest, and secondary forest at blowdowns. When there are abundant ripe legume pods growing along rivers [November–January], titi monkeys are especially common there. They are not found very high up in the trees. They descend to the ground occasionally to drink water, eat insects or fruits, or to cross a swidden.

They live in small troops that number only up to about five or six animals. Sometimes one is found alone. The male always carries the babies. They carry the baby on their back or their belly, and give it to the female to suckle. They call out saying ‘‘ tsokon, tsokon, tsokon, on, on, on ’’ shortly before dawn [around 5 a.m.], later than howler monkeys, and then again later in the morning [around 8 a.m.], and sometimes at midday. The male sings first, and then the rest join in. They sing very loudly, and Matses always hear them from their homes.

Titi monkeys sleep hugging each other on a tree branch among thick vegetation, in vine tangles. Right from where they sleep they sing in the morning. They spread out to eat, but not too far from each other. After eating, they rest at midday. They do not travel far. They are curious, and animals that have not been hunted will stare from the forest edge at people passing by in boats.

Titi monkeys are the only monkeys that eat the ripe mesocarp of cabbage palm [ Euterpe precatoria ] fruits, and that is why they have black teeth [capuchins bite unripe cabbage palm fruits to eat the soft endosperm, but don’t eat the ripe mesocarp]. They eat all types of dicot fruits, especially mannan tsipuis [ Inga spp. and? Pithecellobium (Leguminosae) ] and achu inkuente [ Inga spp. (Leguminosae) ]. Other dicot fruits they eat include: wesnid dëbiate [ Anacardium giganteum . ( Anacardiaceae )], këku [ Couma macrocarpa (Apocynaceae) ], diden këku [ Parahancornia peruviana (Apocynaceae) ], machishte [ Rhigospira quadrangularis and? Mucoa duckei (Apocynaceae) ], okodo mabis [undetermined sp. (Guttiferae)], mamuin [ Rheedia longifolia (Guttiferae) ], chichombid [ Mouriri spp. (Melastomataceae) ], şhannëd [? Brosimum (Moraceae) ], piuşh bëchi [ Helicostylis tomentosa and H. elegans (Moraceae) ], bata [ Pseudolmedia and? Maquira spp. (Moraceae) ], mabis mabiskid [ Chrysophyllum prieurii (Sapotaceae) ], kose [ Manilkara bidentata (Sapotaceae) ], and taëpa [undetermined].

Tables

TABLE 14 Measurements (mm) of Callicebus cupreus from the Yavarí-Ucayali Interfluve

	AMNH		AMNH	FMNH
	Males a 73708♀ 268239♀ 88860♀
HBL	337 (327–346) 5	—	—	315
LT	431 (390–456) 5	—	—	460
HF	92 (90–95) 5	—	93	90
Ear	29 (25–31) 5	—	32	30
CIL	53.4 (49.6–55.8) 9	53.2	52.8	—
OB	35.9 (34.4–37.3) 9	34.5	36.7	37.2
POC	30.1 (27.9–31.9) 9	30.5	31.9	31.9
ZB	40.1 (38.7–42.6) 8	39.1	41.0	41.0
BB	35.4 (33.7–36.7) 9	34.9	35.5	35.1
PPL	26.0 (24.4–28.1) 8	—	26.0	27.0
LMT	18.8 (18.2–19.9) 9	19.2	19.1	19.0
BM1	4.7 (4.4–5.2) 9	4.6	4.3	4.8
M1–M1	20.8 (19.6–22.3) 8	20.9	20.1	21.4
I2–I2	9.9 (9.4–10.2) 7	10.2	10.2	—

a

Summary statistics (mean, observed range in parentheses, and sample size) for measurements of AMNH 73703–73706; FMNH 86989, 86990, 88856, 88858, 88859.