The Taxonomic Status of Mico marcai (Alperin 1993) and Mico manicorensis (van Roosmalen et al. 2000) (Cebidae, Callitrichinae) from Southwestern Brazilian Amazonia Guilherme Siniciato Terra Garbino Received: 03 December 2013 /Accepted: 11 January 2014 # Springer Science+Business Media New York 2014 Abstract Species level taxonomic revisions of New World monkeys are scarce but necessary if we are to obtain a clear picture of this group’ s diversity. In this work I revise taxonomically the little known marmoset Mico marcai¸ originally described as Callithrix argentata marcai Alperin, 1993, in light of more recently collected material. I show that this species is significantly different in facial pigmentation and pelage color from all other marmosets of the genus Mico in the Madeira–Tapajós interfluvium. The form first described as Callithrix manicorensis is treated as a junior synonym of Mico marcai, as there are no morphological differences or geographical barriers between the two putative taxa. Mico marcai occurs in the Aripuanã–Manicoré interfluvium, limited in the north by the Rio Madeira and in the south by enclaves of savannah vegetation. Mico marcai occurs in sympatry with Callibella humilis through at least part of its range, representing the only known case of sympatry of two species of marmoset. The northern range of Mico marcai is relatively undisturbed and has few human settlements, its southern limit, on the other hand, is crossed by a major Brazilian highway, and in the same region at least seven hydroelectric reservoirs are planned to be built. Keywords Callitrichinae . Mico manicorensis . Mico marcai . Taxonomy Introduction The number of recognized New World primate species has more than doubled in the last 30 years, and much of this increase is the result of traditional taxonomic revisions (Groves 2001b). Among callitrichines, a great increase in recognized species was due Int J Primatol DOI 10.1007/s10764-014-9766-4 G. S. T. Garbino (*) Museu de Zoologia da Universidade de São Paulo, Ipiranga 04263-000 São Paulo, Brazil e-mail: [email protected]G. S. T. Garbino Pós-graduação, Museu de Zoologia, Universidade de São Paulo, CEP 04263-000 São Paulo, SP, Brazil
Transcript
The Taxonomic Status of Mico marcai (Alperin1993) andMico manicorensis (van Roosmalen et al.2000) (Cebidae, Callitrichinae) from SouthwesternBrazilian Amazonia
Guilherme Siniciato Terra Garbino
Received: 03 December 2013 /Accepted: 11 January 2014# Springer Science+Business Media New York 2014
Abstract Species level taxonomic revisions of New World monkeys are scarce butnecessary if we are to obtain a clear picture of this group’s diversity. In this work Irevise taxonomically the little known marmoset Mico marcai¸ originally described asCallithrix argentata marcai Alperin, 1993, in light of more recently collected material. Ishow that this species is significantly different in facial pigmentation and pelage colorfrom all other marmosets of the genus Mico in the Madeira–Tapajós interfluvium. Theform first described as Callithrix manicorensis is treated as a junior synonym of Micomarcai, as there are no morphological differences or geographical barriers between thetwo putative taxa. Mico marcai occurs in the Aripuanã–Manicoré interfluvium, limitedin the north by the Rio Madeira and in the south by enclaves of savannah vegetation.Mico marcai occurs in sympatry with Callibella humilis through at least part of itsrange, representing the only known case of sympatry of two species of marmoset. Thenorthern range ofMico marcai is relatively undisturbed and has few human settlements,its southern limit, on the other hand, is crossed by a major Brazilian highway, and in thesame region at least seven hydroelectric reservoirs are planned to be built.
The number of recognized New World primate species has more than doubled in thelast 30 years, and much of this increase is the result of traditional taxonomic revisions(Groves 2001b). Among callitrichines, a great increase in recognized species was due
Int J PrimatolDOI 10.1007/s10764-014-9766-4
G. S. T. Garbino (*)Museu de Zoologia da Universidade de São Paulo, Ipiranga 04263-000 São Paulo, Brazile-mail: [email protected]
G. S. T. GarbinoPós-graduação, Museu de Zoologia, Universidade de São Paulo, CEP 04263-000 São Paulo, SP, Brazil
to the taxonomic works of Rosenberger and Coimbra-Filho (1984) with Leontopithecusand de Vivo (1991) with Callithrix and Mico. These authors were among the first NewWorld monkeys taxonomists to follow the current trend of abandoning the use ofsubspecific category (Groves 2001a, b).
The Amazonian marmoset genus Mico Lesson, 1840, as revalidated by Rylandset al. (2000), comprises 14 recognized species, all of them, with the exception ofM. melanurus, endemic to Brazil (Ferrari et al. 2010; Rylands et al. 2009). Althoughthe taxonomy of the Amazonian marmosets was revised by Hershkovitz in 1977 and deVivo in 1991, seven new species have been described in the last two decades, all ofthem in the Madeira–Tapajós interfluvium, in the southern Brazilian Amazon (Alperin1993; Ferrari and Lopes 1992; Ferrari et al. 2010; Mittermeier et al. 1992; vanRoosmalen et al. 2000; Silva-Júnior and Noronha 1998). In addition, phylogeneticstudies have prompted a reevaluation of the traditional arrangement of just twomarmoset genera: Cebuella, the Amazonian pygmy marmoset, and Callithrix for thelarger marmosets of the Amazon and Brazilian Atlantic forest. The AmazonianCallithrix are now placed in a separate genus Mico Lesson, 1840 (Rylands et al.2000, 2009), and M. van Roosmalen and T. van Roosmalen (2003; see also Aguiarand Lacher Jr. 2003) placed the dwarf marmoset in a new monotypic genus, Callibella.
Of the new marmoset species, Alperin (1993) described Callithrix argentata marcai.The description was based on three individuals, the only known specimens of thetaxon, collected during the Rondon–Roosevelt expedition in 1914. They were previ-ously considered to be Callithrix argentata melanura by Hershkovitz (1977) andCallithrix emiliae by de Vivo (1985).
AlthoughMico marcai has always been considered a valid taxon (Ferrari et al. 2010;Groves 2001a, 2005; Rylands et al. 1995, 2000, 2009; van Roosmalen et al. 1998) thepoorly preserved type material and the absence of additional specimens made thediagnosis and distribution of the species unclear. The type series of Callithrix argentatamarcai was identified as Callithrix emiliae by Silva-Júnior and Noronha (1998) in theirdescription of Callithrix saterei. Moreover, subsequent descriptions of new speciesmade no reference toMico marcai or, when doing so, they overlooked the few availablespecimens and made comparison only with the original text (Ferrari et al. 2010; vanRoosmalen et al. 1998, 2000).
Here I reevaluate the pelage characteristics of Mico marcai and report additionalspecimens beside the type series. In light of material collected more recently, I revisethe distribution and taxonomy of the species.
Methods
Specimens examined are housed in the following Brazilian institutions: InstitutoNacional de Pesquisas da Amazônia (INPA) in Manaus, Museu Nacional do Rio deJaneiro (MN) in Rio de Janeiro, Museu Paraense Emílio Goeldi (MPEG) in Belém,Museu de Zoologia da Universidade de São Paulo (MZUSP) in São Paulo, andUniversidade Federal de Rondônia (UNIR) in Porto Velho.
I examined the three specimens (the holotype, MN 2856, and two paratypes, MN2851, 2867) used in the original description of Callithrix argentata marcai Alperin,1993 to understand properly the morphological characters and individual variation
G.S.T. Garbino
present in the taxon. I then compared the purported diagnostic features to 264 speci-mens (253 skins, 171 crania) of marmosets belonging to the genera Callibella andMicofrom the Madeira–Tapajós interfluvium (Appendix I). The material examined includesthe holotypes of Callibella humilis (van Roosmalen et al. 1998); Mico intermedius(Hershkovitz 1977); Mico nigriceps (Ferrari and Lopes 1992); Mico mauesi(Mittermeier et al. 1992); Mico saterei (Silva-Júnior and Noronha 1998); Micoacariensis (van Roosmalen et al. 2000); Mico manicorensis (van Roosmalen et al.2000) and Mico rondoni Ferrari, Sena, Schneider and Silva-Júnior 2010.
I selected 11 cranial measurements following de Vivo (1991): greatest skull length(GLS), condylobasal length (CBL), zygomatic breadth (ZYB), orbital width (ORB),braincase width (BRW), maxillary toothrow length (MXL), width across upper molars(M–M), width across upper canines (C–C), greatest length of mandible (MAL), lengthof lower postcanine toothrow (MDL), and mandible condylar height (MCH). Theprincipal components analysis (PCA) (no rotation) used the following 10 of the 11variables: GLS, CBL, ZYB, BRW, MXL, M–M, C–C, MAL, MDL, and MCH (Iexcluded BRW as I could not measure it for most individuals). Principal components(PC) with eigenvalues greater than 1 were kept and PC scores were plotted to showgrouping in the morphospace.
Only adult specimens, with fused basioccipital–basisphenoid suture and eruptedpermanent dentition, were included in the analysis. Adult male and female specimenswere lumped, as I assumed sexual dimorphism was not significant (Ford 1994; Marroigand Cheverud 2009). From the 174 skulls (the three fromMico marcai type series plus171) I selected, based on age class and integrity, and measured 113 belonging toCallibella andMico from the Madeira–Tapajós interfluvium and included in PCA onlythe untufted neighbors of Mico marcai in whose individuals all 10 variables weremeasured.
Results
Morphometrics
The loading factors of the 10 cranial variables and the contribution of the first twocomponents in the PCA are shown in Table I: Principal component 1 (PC1) wasresponsible for 57.9 % of the variation, while PC2 was responsible for 19.9 %.Together the two components were responsible for 77.9 % of the variation.
The loading factors of PC1 show that it is related to size differences, while PC2expresses differences in shape. However, the specimens, which included only theuntufted geographic neighbors of Mico marcai, showed much overlap among species(Fig. 1).
Qualitative Craniodental Characters
The nasal profile of the skull is straight (not convex as in Callithrix), a distinctivefeature of all the Amazonian marmosets, i.e. Callibella, Cebuella, and Mico. Thezygomatic arch is large on the orbital margin; a posterior nasal spine is absent; and awell developed petrosal spine is present (Fig. 2). The dental formula of Mico marcai is
Taxonomic Status of Mico marcai and Mico manicorensis
typical of callitrichines (Hershkovitz 1977): I 2/2, C 1/1, P 3/3, M 2/2. The outer upperincisors have a higher crown than the inner ones and the tips diverge from a central axisin centric view, which results in the interdigitated occlusional pattern typical ofmarmosets; P2 lacks a protocone; the P4 is waisted (Kay 1994). Both upper molarslack a hypocone. The entocingulum of M1 completely wraps the protocone, a distinc-tive character of Callibella, Cebuella, and Mico but not Callithrix (de Vivo 1991).
A mesiostyle and a distostyle are present on the i1 and i2. The i1,2 are high crowned,being subequal in crown height to the canines; the incisors and the canines are
Table I Scores and contributions of each cranial variable in the first two principal components of a principalcomponent analysis of Mico marcai and its untufted neighbor species in the Madeira–Tapajós interfluvium
Variable Principal component 1 Principal component 2
Greatest skull length 0.881 0.684
Condylobasal length 0.994 –0.174
Zygomatic breadth 0.082 0.122
Braincase width 0.208 0.184
Maxillary toothrow length 0.395 –0.320
Width across upper molars 1.225 –0.411
Width across upper canines 0.150 –0.228
Greatest length of mandible 0.532 –0.568
Length of lower postcanine tooth row 0.729 0.248
Mandible condylar height. 0.535 0.575
Variation explained 57.9 % 19.9 %
Initial eigenvalues 5.595 1.584
Fig. 1 Plots of scores for the first two principal components from principal component analysis of cranialvariables in Mico marcai and its untufted neighbor species in the Madeira–Tapajós interfluvium. Crosses =Mico rondoni; open losangles = M. marcai; open triangles = M. saterei; starburst = M. nigriceps.
G.S.T. Garbino
staggered, giving a “V” shape to the anteriormost portion of the mandible (Hershkovitz1977). These teeth, however, are not as staggered as in Callithrix (Hershkovitz 1975).The coronoid has a convex tip and is nearly at the same level as the condylar process.The angular process projects ventrally to the horizontal ramus and posteriorly to thevertical ramus of the mandible (Fig. 2). The condylar process is located slightly abovethe dental plane, a common feature in Mico species.
External Morphology
Comparison of the chromogenetic fields (sensu Hershkovitz 1968) showed that thethree specimens from the type series of Mico marcai (MN 2851, 2856, 2857), and twoadditional ones including the holotype of M. manicorensis (INPA 2511), may begrouped together and diagnosed by the characters mentioned below. The only differ-ence in the coloration between two additional specimens and the type material of Micomarcai is that the former have more vivid colors than the latter. This is probablybecause the specimens in the MN (the type series) were collected in 1914, whereas thespecimen fromMPEG (MPEG 22960) was collected in 1993 and the one from INPA in1997, and the older specimens may be slightly faded.
Mico marcai exhibits a pigmented face except in the region around the nostrils andbetween the eyes. A vertical pigmented stripe is present between the nostrils. The ears
Fig. 2 Skull of Mico marcai holotype (MN 2856) in (from above to the bottom): dorsal, ventral, and lateralviews of the skull and mandible in lateral profile. Scale bar = 10 mm.
Taxonomic Status of Mico marcai and Mico manicorensis
are unpigmented and mottled with melanin. The crown is dark chestnut-brown, contrast-ing with the dark gray nape and mantle. Hairs on the nape and mantle region present asmall light gray basal band, followed by a larger plumbeous gray middle one and a silverygray apical band. The saddle and rump have the same banding pattern, which makes thedorsum an overall light gray mottled with dark gray. Although the mantle is not welldefined, the overall color in this region is lighter than in the region posterior to it. There is awhite triangular patch in the center of the forehead that is part of a circular white rimextending to the sides of the forehead and to the face, up to the region posterior to the eyes.The forelimbs are light brown with some silvery gray hairs on the proximal portion. Thehind limbs are reddish-brown to ochraceous; the hair in this region has a clear buff basalband and a reddish-brown terminal band. The ventral region is silvery-white on the forelimbs and gular, neck, and anterior chest regions; posterior to this point, the overall color isreddish-brown to ocher, similar to that found on the hind limbs. The tail is completelyblack in the five specimens examined, except for the basal portion in the specimens MN2851, 2856, and 2857, which have reddish-brown ochraceous hairs.
The specimen INPA 2511 (Fig. 3), here identified asMico marcai, is the holotype ofCallithrix manicorensis (now Mico manicorensis), described as a new species by vanRoosmalen et al. (2000). It is a juvenile female still with milk teeth along withpermanent first molars and unerupted second molars. Based on the principle of priorityof the International Code of Zoological nomenclature (ICZN, 1999), Callithrixmanicorensis is thus a junior synonym of Callithrix argentata marcai.
The specimen MPEG 22960, from “Humaitá-Apuí Road (BR-230), km 292, leftmargin of rio Aripuanã” is nearly a topotype of Mico marcai and differs from theanalyzed sample only in exhibiting a slightly darker dorsal pelage.
Another two specimens (INPA 2512, 3930), both adults with tooth wear and constitutedonly by skulls and partial skeletons, were also identified as Callithrix manicorensis by vanRoosmalen et al. (2000). Figure 5 in van Roosmalen et al. (2000) shows them alive, but theirskinswere not preserved.Owing to the reported collecting locality of these two specimens andto the fact that van Roosmalen et al. (2000) mention that both individuals exhibited the samepelage characters as the INPA2511 specimen, they can be assigned toMicomarcai. However,a definitive identification of these individuals is impossible because the skins were lost.
Comparison with Similar and Neighboring Taxa
The dwarf marmoset Callibella humilis can be differentiated from Mico marcai by itssmaller size, being almost a third that ofM. marcai (Table II). The pelage is also distinctive:Callibella humilis has a white patch on the brow that extends to the region behind the eyes;in Mico marcai the white patch is more pronounced on the forehead, being only faintlydistinguishable in the region behind the eyes. The dwarf marmoset also has a more hirsuteear, with a small tuft of hair emerging from the lower auricular lobe.Callibella humilis doesnot have ochraceous hind limbs and its tail is not uniformly black but consists ofmultibanded hairs that may be disposed in rings, as in specimen MPEG 24769.
Mico chrysoleucos is the species most different fromM. marcai in pelage characters,having an overall golden-white pelage, a hairy tufted ear, and a ringed tail.
Mico intermedius has a clearly bizonal pelage, with the anterior portion white andthe posterior mottled irregularly with brown (de Vivo 1991).Mico intermedius also hasgolden-yellow hip patches extending along the thighs.
Taxonomic Status of Mico marcai and Mico manicorensis
TableII
Com
parisonof
cranialm
easurements,inmillim
eters,with
themeanfollowed
by±standard
deviationandsamplesize
(N)inparentheses,of
Micomarcaiand
othermarmoset
speciesfrom
theMadeira–T
apajós
interfluvium
Taxon
Greatest
skulllength
Condylobasal
length
Zygom
atic
breadth
Orbitalwidth
Braincase
width
Maxillary
toothrow
length
Width
across
uppermolars
Width
across
uppercanines
Greatestlength
ofmandible
Lengthof
lower
postcanine
toothrow
Mandible
condylar
height
Callib
ella
humilis
36.8
±1.16
(3)
29.8±0.91
(3)
24.7
±1.12
(3)
21.9±0.38
(2)21.2±0.80
(3)
9.5±0.28
(3)
12.0
±0.72
(3)
9.0(3)
22.9
±0.26
(3)
8.7±0.67
(2)
12.2
±0.29
(3)
Micomarcai
45.2
±1.16
(6)
36.6±1.01
(5)
30.1
±0.89
(6)
25.6±0.34
(4)25.1±0.70
(5)
11.3
±0.22
(6)
15.0
±0.53
(6)
11.6
±0.24
(6)
29.6
±1.13
(6)
10.8
±1.16
(6)
14.8
±0.59
(6)
M.a
cariensis
48.0
(1)
38.5
(1)
30.6
(1)
27.0
(1)
26.6
(1)
11.8
(1)
15.9
(1)
12.3
(1)
31.9
(1)
10.7
(1)
14.2
(1)
M.chrysoleucos
46.5±0.92
(29)
36.1
(1)
31.2±1.18
(27)
—24.8
(1)
11.7±0.25
(25)
15.7±0.45
(26)
11.9±0.67
(24)
30.6±1.14
(29)
10.6±0.27
(24)
14.8±0.75
(26)
M.h
umeralifer
45.2±1.08
(22)
—29.6±1.38
(20)
——
11.2±0.32
(24)
15.2±0.46
(23)
11.5±0.56
(21)
28.8±1.40
(23)
10.2±0.61
(23)
13.6±0.70
(24)
M.intermedius
47.6
(1)
36.9
(1)
——
25.9
(1)
11.4
(1)
15.8
(1)
11.6
(1)
30.7
(2)
10.3
(2)
14.3
(2)
M.m
auesi
46.6
±1.22
(4)
36.9±2.65
(2)
29.8
±2.27
(4)
25.8
(1)
26.5±1.28
(2)
11.9
±0.05
(4)
15.7
±0.46
(4)
11.6
±0.87
(4)
30.5
±2.64
(5)
11.1
±0.30
(5)
14.5
±1.92
(5)
M.m
elanurus
46.3±1.13
(18)
—30.3±1.04
(16)
——
11.2±0.29
(22)
15.0±0.56
(23)
11.2±0.39
(21)
29.9±0.92
(19)
10.2±0.36
(21)
14.5±0.87
(21)
M.n
igriceps
46.0
±0.88
(5)
37.0±1.02
(2)
39.8
±0.61
(3)
—26.2±0.36
(2)
11.6
±0.38
(5)
15.7
±0.32
(5)
12.0
±0.43
(5)
30.8
±0.63
(5)
10.9
±0.34
(5)
14.6
±0.54
(5)
M.rondoni
45.5
±0.55
(5)
35.8±0.82
(4)
29.7
±0.93
(5)
—25.8±0.38
(4)
11.6
±0.30
(5)
15.4
±0.32
(5)
11.3
±0.41
(5)
29.4
±1.25
(5)
10.6
±0.11
(5)
14.8
±0.72
(5)
M.saterei
46.5
±0.86
(7)
37.3±0.57
(7)
30.9
±1.08
(7)
26.8±0.68
(3)26.4±0.63
(7)
11.7
±0.37
(7)
15.8
±0.34
(7)
11.6
±0.91
(6)
30.0
±1.32
(7)
10.9
±0.30
(7)
14.7
±1.29
(7)
Micosp.
46.4
(1)
——
——
11.0
(1)
15.6
(1)
12.0
(1)
31.4
(1)
10.3
(1)
14.7
(1)
G.S.T. Garbino
Mico nigriceps presents an overall darker coloration pattern and pigmented ears. Theforelimbs are darker on the manus, lightening gradually more proximally to the body. Italso exhibits hip patches extending to the thighs, which are absent inMico marcai. Bothtaxa have a conspicuous white patch in the center of the forehead.
The species most similar to Mico marcai in pelage characters are M. nigriceps andM. rondoni. Mico marcai differs from M. rondoni in having a more pigmented face(unpigmented only around nostrils), with hands the same color as the arms (not darkerthan the arms, as in M. rondoni), not showing a light gray anterior dorsum, and havingmore ochraceous hind limbs (Fig. 4).
The individual referred to here asMico sp. (MPEG 22955) differs fromM. marcai inhaving a less pigmented face and having hip patches. More specimens are needed,however, to classify this taxon correctly.
Systematics
Mico marcai (Alperin, 1993)
Marca’s Marmoset
Callithrix argentata melanura: Hershkovitz (1977, p. 569). Part, not Simia melanurusGeoffroy in Humboldt (1812).
Callithrix emiliae: de Vivo (1985, p. 105). Part, not Hapale emiliae Thomas, 1920.de Vivo (1991, p. 29). Part, not Hapale emiliae Thomas, 1920. Silva-Júnior andNoronha (1998, p. 27). Part, not Hapale emiliae Thomas, 1920.
Callithrix argentata marcai Alperin (1993, p. 325). Type locality “Foz do rioCastanho (= Roosevelt), afluente esquerda do rio Aripuanã, Estado do Amazonas,Brasil.”
Fig. 4 Dorsal view of (a)Mico rondoni (voucher number MZUSP 20142) and (b)Mico nigriceps (MPEG 21998).
Taxonomic Status of Mico marcai and Mico manicorensis
Callithrix marcai: Rylands et al. (1995, p. 116); van Roosmalen et al. (1998, p. 3).Name combination: van Roosmalen et al. (2000, p. 4).
Callithrix manicorensis van Roosmalen et al. (2000, p. 3). Type locality “SeringalSão Luis, east bank of the middle Rio Madeira, in the vicinity of the town of Manicoré,state of Amazonas, Brazil.”
Mico marcai: Rylands et al. (2000, p. 73). First use of current name combination.Alperin (2002, p. 126); Marroig and Cheverud (2009, p. 233); Rylands et al. (2009,p. 27; 2012, p. 2); Rylands and Mittermeier (2009, p. 27; 2013, p. 311); Ferrari et al.(2010, p. 701; Fig. 4); Ennes et al. (2013, p. 327); Garbino et al. (2013, p. 331).
Mico manicorensis: Rylands et al. (2000, p. 73). Name combination. Aguiar andLacher Jr. (2009, p. 359); Alperin (2002, p. 127); Ferrari et al. (2010, p. 705); Ford andDavis (2009, p. 414; Table I); Garbino et al. (2013, p. 331); Rylands and Mittermeier(2009, p. 27; 2013, p. 312); Rylands et al. (2009, p. 27; 2012, p. 2); van Roosmalen andvan Roosmalen (2003, p. 4); Veracini (2009, p. 234).
Callithrix (Mico) marcai: Groves (2001a, p. 132). Name combination. Groves(2005, p. 131).
Callithrix (Mico) manicorensis: Groves (2005, p. 131). Name combination.
Holotype
The type, designated by Alperin (1993, p. 325) is specimenMN 2856, collector number31, male, adult with fused basilar suture. Collected at the type locality byComissão Rondon members on April 9, 1914. Open skin, skull. The right tibiais still in the skin. Deposited in the collection of the Museu Nacional do Rio deJaneiro, Brazil.
Paratypes
MN 2851, collector number 33, unknown sex, adult with fused basilar suture, collectedon April 10, 1914. Skin, skull. This is the best preserved specimen of the type series,because the skin is not open. MN 2857, unknown sex, adult, collected on April 8, 1914.Open skin, skull. Both hind limb bones are still in the skin. The specimens werecollected by Comissão Rondon at the type locality and are deposited in the sameinstitution as the holotype.
Type Locality
Foz do rio Castanho (= Rio Roosevelt), left margin affluent of the Rio Aripuanã,municipality of Novo Aripuanã, state of Amazonas, Brazil (07°33′S 60°43′W). Theholotype still bears the original Comissão Rondon label, with the locality “Foz doCastanhas” hand written on it. Alperin (1993) used the variant Rio Castanho. Althoughnot doubting that the type locality was near the mouth of the Rio Roosevelt, Rylandset al. (2000, p. 74) argued that the Rio Castanho and Rio Roosevelt may not besynonyms, and that the former might refer to an “Igarapé Castanho,” a smallaffluent of the Rio Aripuanã, just below the mouth of the Roosevelt. Alperin(2002) recorded that there was also a small village near the mouth of the RioRoosevelt (left bank) called Castanho. This would explain the locality being
G.S.T. Garbino
described by Hershkovitz (1977; locality 197b, p. 933) as “Castanho, RioCastanhas (= Rio Roosevelt).” Alperin (2002) refuted the suggestion ofRylands et al. using compelling evidence that Castanho and Castanhas referto the Rio Roosevelt.
Distribution
The updated known records of Mico marcai, and specimen-based records of theneighboring species are shown in Fig. 5 and listed in Appendix II. I also mappedspecimen MPEG 22955 (indicated with an asterisk) from the east margin of the Rio dosMarmelos (mentioned in Rylands et al. 2009), the characters of which do not fitany known taxon of Mico and is herein referred as Mico sp. (Appendix II).
The right bank of the Rio Madeira limits the northern distribution of Mico marcai(Fig. 5). The species is limited eastwards by the middle and lower portions of the RioAripuanã, and the Rio Manicoré serves as the western limit to its distribution (Fig. 5).The southern limit of the distribution of Marca’s marmoset probably lies in the regionof the lower and middle Rio Roosevelt. In this region, between the Rios Ji-Paraná andRoosevelt there are enclaves of savannah vegetation that would restrict the species’dispersal southwards (Ferrari 1993).
The range of Mico marcai encompasses that of Callibella humilis (Garbino et al.2013; Rylands et al. 2008, 2009; van Roosmalen et al. 1998) and this is the only
Fig. 5 Known locality records, based on museum voucher specimens ofMico marcai (●) and its type locality(★). Localities of the neighboring species Mico chrysoleucos (▲), Callibella humilis (+), Mico intermedius(♦), Mico nigriceps (■), and Mico sp. (❋) are also plotted on the map. The shaded areas represent opensavannah vegetation patches. Dashed lines indicate Brazilian federal state boundaries. For the localitiescoordinates see Appendix II.
Taxonomic Status of Mico marcai and Mico manicorensis
known case where two marmoset species are sympatric. Differences in sizeobserved between the two morphologically similar species (Fig. 1) surely reflectniche-partitioning related to sympatry (Heymann 1997) and both may even formmixed species groups.
Diagnosis
Mico marcai is distinguished by a conspicuous white patch in the center of theforehead, a dark crown contrasting with the immediately posterior region, indistinctmantle, hands with same color as the arms, overall dorsal coloration light silvery graymottled with dark plumbeous gray, reddish-brown to ochraceous hind limbs, and ablack tail with an ochraceous basal ring (Fig. 3). The otherMico species from the sameregion do not differ significantly in cranial measurements from Mico marcai or fromeach other (Tables I and II).
Ontogenetic Variation
It may be argued, since the holotype ofMico manicorensis is a juvenile, that the pelagetraits cannot be compared to the three specimens from the type series of Mico marcai.However, I have verified that significant alterations, especially in the crown pattern andthe saddle and rump, occur only in tufted species of marmosets, as can be confirmed bythe following specimens: Callibella humilis (based on van Roosmalen et al. 1998),Mico chrysoleucos (specimen INPA 4110), M. humeralifer (MPEG 9213),M. intermedius (MPEG 12599), and M. saterei (MPEG 23960, INPA 5672); this lastis an untufted species that may be more closely related to tufted forms (Schneider et al.2012; Sena et al. 2002; Silva-Júnior and Noronha 1998). Mico marcai is a member ofthe untufted group of Amazonian marmosets, the juveniles of which show the samebasic pelage pattern as the adults, albeit exhibiting more hairy ears, as I have confirmedfor M. argentatus (specimens MPEG 165, MZUSP 11367), M. leucippe (MN 4798),M. nigriceps (MPEG 22956), and M. rondoni (MPEG 21892).
Conservation
The last assessment of the International Union for the Conservation of Nature and NaturalResources (IUCN) red list, categorized Mico marcai as “data deficient,” whileM. manicorensis was classified in the conservation status of “least concern” (Rylandsand Silva-Júnior 2008a, b). The data presented in this paper will hopefully remove Micomarcai from the “data deficient” category, as it is now confirmed to occur in a relativelyundisturbed area, near the right margin of the RioMadeira. Although there are at present nolegally protected areas encompassing the area of occurrence of this species, the northernrange of Mico marcai is relatively well protected (Rylands and Silva-Júnior 2008a).
The Brazilian government, however, plans to build at least seven hydroelectric damsin the Aripuanã basin, along the Rios Roosevelt and Aripuanã (Farias 2012). Permanentflooding caused by the reservoirs will undoubtedly shrink the suitable habitat of Micomarcai. Recently, Ennes et al. (2013) reported the rediscovery of Mico marcai aroundits type locality and alerted the scientific community of the coming threats to the habitatof the species there. Moreover, the southern limit of the distribution of Mico marcai is
G.S.T. Garbino
crossed by the Brazilian highway BR-230, the Transamazônica, which favor humansettling in the area (Fearnside 1986).
These factors will surely threat not only Marca’s marmoset but also other animal andplant species in this rich and yet unknown area, which includes an undescribed form oftiti-monkey (Callicebus).
Conclusion
There are now 13 recognized species in the genus Mico (Rylands et al. 2009) and theform from the right bank of the Rio dos Marmelos may be a new species. The range ofMico marcai is defined to a restricted area between the rivers Manicoré and lower andmiddle Aripuanã and lower Roosevelt. It is consequently the only Mico that occurs insympatry with Callibella humilis. This work reinforces the importance of species-level,morphology-based taxonomy, and the underlying need of more studies involvingmarmoset taxonomy and systematics.
Acknowledgments I thank Mario de Vivo, Vitor Piacentini, Carla de Aquino, Rafael Marcondes, GabiRezende, and Marcelo Weksler for the critical revision of early versions of the manuscript and Joanna Setchelland three anonymous reviewers, whose comments and suggestions were most helpful. I thank Felipe EnnesSilva for sharing valuable information regarding Mico marcai. I thank Victor Fonseca Silva for the MPEGphotos and information. I also thank the curators and staff of the collections I visited: José de Souza e Silva-Júnior (Museu Paraense Emílio Goeldi), João A. Oliveira and Sérgio M. Vaz (Museu Nacional), Mario deVivo and Juliana Gualda (Museu de Zoologia da USP), Maria Nazareth Ferreira da Silva and Manoela Borges(Instituto Nacional de Pesquisas da Amazônia), and Mariluce Messias (Universidade Federal de Rondônia).This work was funded by CAPES.
Appendix I
The skins and skulls examined are deposited in the following Brazilian collections,listed in alphabetical order by their acronyms: Instituto Nacional de Pesquisas daAmazônia (INPA) in Manaus, Museu Nacional do Rio de Janeiro (MN) in Rio deJaneiro, Museu Paraense Emílio Goeldi (MPEG) in Belém, Museu de Zoologia daUniversidade de São Paulo (MZUSP) in São Paulo, and Universidade Federal deRondônia (UNIR) in Porto Velho.
Callibella humilis (3): BRAZIL: Amazonas: Left margin of lower Rio Aripuanã,Nova Olinda settlement, municipality Novo Aripuanã (INPA 4090, MPEG24769); Rio Atininga, right margin, 10 km SE from the town of Manicoré(INPA 4091).
Mico acariensis (1): BRAZIL: Amazonas: Right margin of lower rio Acarí, next tothe confluence with the rios Sucunduri and Canumã (INPA 3931, holotype).
Taxonomic Status of Mico marcai and Mico manicorensis
5022, 5028, 5029, 5030, 11409, 11410, 11411); Prainha, right margin of Rio Aripuanã(MZUSP 13466, 13467); Nova Olinda do Norte (INPA 4039); Rio Madeira, 5 km southof the mouth of Paraná do Canhumã, municipality of Nova Olinda do Norte(INPA 4110).
Mico intermedius (5): BRAZIL: Amazonas: Left bank of rio Aripuanã, mouth of rioGuariba (MN 8156, holotype); Rio Roosevelt, right bank, close to rio Aripuanã (MPEG23065); “Trilha do Roosevelt”, Pousada Rio Roosevelt, right bank of rio Roosevelt,municipality of Novo Aripuanã (MZUSP 35320); Mato Grosso: Aripuanã (MN 2850,MPEG 12599, paratype).
Mico marcai (7): BRAZIL: Amazonas: Humaitá-Apuí Road (BR-230), km 292, lefttmargin of Rio Aripuanã (MPEG 22960); Mouth of rio Castanhas (= rio Roosevelt)(MN 2851, 2867, paratypes, MN 2856, holotype); Santa Cruz settlement, right marginof Rio Mariepauá (INPA 2512); Seringal São Luiz, margem direita do Rio Madeira,5 km S of the town of Manicoré (INPA 2511, holotype of Callithrix manicorensis;INPA 3930).
Mico mauesi (9): born in captivity, Manaus (INPA 4107); BRAZIL: Amazonas: RioMaués-Açu, left bank, municipality of Maués (INPA 4082, 4105, 4109; MPEG22177, holotype); Rio Maués-Maraú, left bank (INPA 4106); Santa Maria, rightbank of Rio Abacaxis, municipality of Nova Olinda do Norte (MPEG 23962,23963, 23964).
Mico melanurus (48): locality unknown (MN 2849, 2858, 2859; MPEG 13295);BRAZIL: Mato Grosso: Cáceres (MN 5845, 5847, 5849); Cidade laboratórioHumboldt, Aripuanã (MPEG 13289, 13290, 21395, 21396, 15266, 15267); Mata doToscano, municipality of Cáceres (MN 2848, 2852); Rio Aricá (MN 2854, 2855;MZUSP 6327, 6328, 6330, 6333, 11415); Salto, Rio Jaurú (MN 2853); SantoAntonio do Leverger (MZUSP 4263, 4264, 4265, 4266); Wood close to “Casa dePedra”, Chapada dos Guimarães (MN 24912, 24913); Mato Grosso do Sul: Corumbá(MZUSP 3367, 3368, 3369, 3370, 3376, 3377, 7909. 7910, 7911, 24734); FazendaPanatanlzinho, Posto Espiridão (MN 25008); Palmeiras (MZUSP 6329, 6332);
G.S.T. Garbino
Rondônia: BR-429 road, municipality of São Francisco doGuaporé (SFF97/01, SFF97/03);UHE Rondon II, municipality of Pimenta Bueno (UFRO- M192, M346, M356).
Mico nigriceps (10): BRAZIL: Amazonas: Humaitá-Apuí Road (BR-230), km 16(MPEG 21999, paratype); Humaitá-Apuí Road (BR-230), km 17 (MPEG 21996,paratype; MPEG 21998, holotype); Lago dos Reis, Humaitá-Apuí Road (BR-230),right bank of Rio Madeira (MPEG 21956, 21957, 21958, 21959); Tenharinsettlement, left bank of Rio dos Marmelos, Humaitá-Apuí Road (BR-230), km 104(MPEG 22961, 22962); Rondônia: Calama, right bank of rio Ji-Paraná (MPEG 21997,paratype).
Mico rondoni (17): locality unknown (UFRO-M396); BRAZIL: Rondônia: BR-364road, municipality of Porto Velho (UFRO-M078); Campus of the Universidade Federalde Rondônia, municipality of Porto Velho (UFRO-M048); Garimpo Bom Futuro,municipality of Ariquemes (MN 66399); Mouth of Rio Urupá, Gi-Paraná settlement,vila Rondônia (MN 23827); Porto Velho (UFRO-M162, M165); Santa Bárbara(MZUSP 20142); UHE Samuel, Rio Jamari, municipality of Porto Velho (MN29476, paratype; MPEG 21888, MPEG 21366, 21647, 21660, 21892, 21896,paratypes, MPEG 21893, holotype).
Mico saterei (13): unknown locality (INPA 4104, MPEG 23961); BRAZIL:Amazonas: Mouth of Rio Canumã, right bank, Municipality of Borba (MPEG23955, holotype, MPEG 23956); São João, left bank of Rio Marimari, munic-ipality of Borba (MPEG 23957, 23958); Terra Preta, right bank of RioAbacaxis, municipality of Nova Olinda do Norte (MPEG 23959, 23960); RioMari mari, Maués Indian reserve (INPA 41010); Rio Mari mari, Vila Batista(INPA 4081, 4102, 4103, 4040, 5672).
Mico sp. (1): BRAZIL: Amazonas: Opposite of Tenharin settlement, right bank ofRio dos Marmelos, Humaitá-Apuí Road (BR-230) (MPEG 22955).
Appendix II
Gazetteer of localities plotted on the map of Fig. 5.
1. Pousada Rio Roosevelt, “Trilha do Roosevelt,” right bank of Rio Roosevelt,municipality of Novo Aripuanã (08°32'14'' S 60°54′29′′W).
2. Rio Aripuanã, right bank, close to mouth of Igarapé Maracanãzinho (08°30′S59°30′W) (estimated).
3. Calama, right bank of Rio Ji-Paraná (08°03′S 62°53′W).4. Tenharin settlement, left bank of Rio dos Marmelos, Humaitá-Apuí Road
(BR-230), km 104 (07°57′S 62°03′W).5. Opposite of Tenharin settlement, right bank of Rio dos Marmelos, Humaitá-Apuí
Road (BR-230) (07°57′S 62°03′W).6. Left bank of Rio Aripuanã, mouth of Rio Guariba (07°43′S 60°35′W).7. Rio Roosevelt, right bank, close to Rio Aripuanã (07°36′S 60°42′W).8. Mouth of Rio Castanho (variants Castanhas, Castanha, Castanhos) (= Rio
Roosevelt) (07°33′S 60°43′W). Type locality of Mico marcai.9. Humaitá-Apuí Road (BR-230), km 292, left margin of Rio Aripuanã (07°32′S
60°40′W).
Taxonomic Status of Mico marcai and Mico manicorensis
10. Lago dos Reis, Humaitá-Apuí Road (BR-230), right bank of Rio Madeira (07°31′S 62°52′W).
11. Humaitá-Apuí Road (BR-230), km 306, right margin of Rio Aripuanã (07°30′S60°34′) (estimated).
12. Humaitá-Apuí Road (BR-230), km 16, km 17(07°29′S 62°56′W) (estimated).13. Prainha, right margin of Rio Aripuanã (07°16′S 60°22′W).14. Rio Atininga, right margin, 10 km SE from the town ofManicoré (05°54′S 61°15′W).15. Seringal São Luiz, margem direita do Rio Madeira, 5 km S of the town of
Manicoré (05°50′28′′S 61°18′19′′W). Type locality of Callithrix manicorensis.16. Santa Cruz settlement, right margin of Rio Mariepauá (05°49′S 60°32′W).17. Left margin of lower Rio Aripuanã, Nova Olinda settlement, municipality Novo
Aripuanã (05°30′63′′S 60 24′61′′W).18. Rio Madeira, 5 km south of the mouth of Paraná do Canhumã, municipality of
Nova Olinda do Norte (04°00′S 59°15′W).19. Lago Curupira, norte do Cana (03°46' S 58°32′W).20. Lago do Batista (03°20'S 57°53′W).
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