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Ting, Suyin, Wang, Xiaoming, Meng, Jin (2023): Cranial And Postcranial Morphology Of The Insectivoran-Grade Mammals Hsiangolestes And Naranius (Mammalia, Eutheria) With Analyses Of Their Phylogenetic Relationships. Bulletin of the American Museum of Natural History 2023 (463): 1-129, DOI: 10.1206/0003-0090.463.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-463/issue-1/0003-0090.463.1.1/Cranial-and-Postcranial-Morphology-of-the-Insectivoran-Grade-Mammals-Hsiangolestes/10.1206/0003-0090.463.1.1.full

Phylogenetic relationships among sloths (Folivora) have been extensively studied in the past few decades using maximum parsimony approaches. Recently, Bayesian phylogenetic methods also began to be employed for this task, with advances in methods for data partitioning and tip-dating analyses leading to exciting new possibilities in morphological phylogenetics. In this context, we assembled the largest morphological data set ever applied to sloths and reassessed their phylogeny and divergence times, evaluating alternative models of partitioning and dating in a Bayesian framework. The updated phylogeny of sloths is largely in agreement with previous morphological studies, with Bradypus recovered as sister to Eufolivora, the presence of two major sloth clades (Mylodontoidea and Megatherioidea) and Choloepus among Megalonychidae. However, the present study yields some important advances in understanding the relationships of genera with historically unresolved or controversial allocations. The major sloth clades diversified from the Late Eocene to the Early Miocene. Homoplasy-based partition models outperformed anatomical partitioning and unpartitioned analyses, with considerable impacts on topology and posterior probabilities. Estimates obtained using homoplasy-partitioned models with Bayesian analyses were in strong agreement with those of maximum parsimony. We emphasize the importance of model comparison with Bayes factors and the assessment of synapomorphies in Bayesian morphological phylogenetics.

Prepotheriinae PP = 99, age = 17.11 Mya (16.20–18.61). The clade is composed of the genera Prepotherium, Planops and Prepoplanops and was consistently recovered in all of our analyses. Planops was recovered as the sister taxon to a clade uniting the other two genera in all but four analyses, those with H models— IW10_e, IW10_p, IW5_e and IW5_p. In these cases, Prepotherium assumed a stem position. Prepotheriinae was supported by 14 synapomorphies (ten for both methods and four exclusively for BI): absence of fossa anterior to Cf1 (BI); posterior segments of temporal lines run anterior to but closely parallel the nuchal crest (BI); postorbital process roughly at the level of maxillary foramen; triangular occiput in posterior view (BI); condyles lie just posterior, almost continuous with hypoglossal foramina (BI); absence of styliform process of ectotympanic; anteroposterior orientation of entotympanic; ventrolateral orientation of stylohyal articulation; presence of glenoid posterior shelf; strongly marked brachiocephalicus crest of humerus; medial epicondyle of humerus rounded to slightly pointed laterally; proximal half of femur wider than distal half; calcaneus tuberous and expanded, with distal apex rounded; and confluence of sustentacular facet and cuboid surface of calcaneus.

Megalonychinae PP = 100, age = 10.82 Mya (8.60–13.21). This clade is composed of the genera Megalonyx (Megalonychini) and Pliometanastes, plus a clade uniting Pliomorphus, Antillean megalonychines and Choloepus (Choloepodini) and a clade of intertropical megalonychids (Ahytheriini). Alternative arrangements were observed for the relationships among Megalonyx, Pliometanastes and the other megalonychines: as successive sister taxa, diverging earlier than all other megalonychines— Megalonyx earliest (H models EW and IW100, par_EW and par_ IW100), Pliometanastes earliest (UN and A models, especially those including ACRV); Megalonyx and Pliometanastes as sister taxa to each other, and this clade sister to all other megalonychines (A models with equal rates, excepting A1_e); or as sister to Ahytheriini (H models IW10 and IW5, par_IW10 and par_IW5). Megalonychinae was supported by 59 synapomorphies (24 for both methods and 35 exclusively for BI): Cf1 and cf1 strongly depressed relative to molariforms; Cf1 smaller than smallest molariform (BI); long axis of molariforms oblique to toothrow along its entire length; trapezoidal Mf2 and Mf3 cross-section; trapezoidal mf2 cross-section; trigonal Mf4 cross-section; mandibular horizontal ramus length less than two times its depth (BI); ascending ramus of mandible completely covers the posterior teeth in lateral view (BI); three processes of ascending mandibular ramus equidistant (BI); absence of a medial ridge running along anterior edge of coronoid process (BI); maximum height of coronoid process shorter than its anteroposterior length (BI); intermediate development of angular process; symphysial spout much shorter than the length of molariform toothrow (BI); posterior external opening of mandibular canal opens laterally on horizontal ramus (BI); low and broad braincase (BI); reflexed basicranial/basifacial axis; nuchal crest continuous with dorsal edge of zygomatic process of squamosal (BI); premolariform portion of palate roughly equal to, or longer than molariform toothrow (BI); presence of fossa at the anterior edge of maxilla, lateral to external nares (BI); presence of ventral extension in maxilla for dental alveoli (BI); ventral extension in maxilla for dental alveoli only posteriorly (BI); jugal does not participate in rim of maxillary foramen (BI); absence of dorsal process of premaxilla; presence of a crest at median suture of palatine (BI); presence of alisphenoid-parietal contact (BI); anteroventrally inclined zygomatic process of squamosal; absence of a marked postorbital constriction (BI); postorbital process well anterior to maxillary foramen; semicircular occiput, in posterior view (BI); hypoglossal foramen smaller than jugular foramen (BI); basioccipital narrow and convex mediolaterally (BI); median fusion of posterior alae of vomer; ectotympanic fused dorsally; carotid foramen fully exposed in ventral view (BI); anteroposteriorly oriented entotympanic (BI); dorsal edge of entotympanic with a strong concave curvature in lateral view, with dorsal projection at anterior end (BI); lateral process of entotympanic extending above anterior portion of tympanic, forming a portion of the roof of tympanic cavity (BI); bulbous, mediolaterally expanded paroccipital process; absence of paroccipital process foramen (BI); nuchal and exoccipital crests diverge proximally and converge distally (BI); posteroventrolateral stylomastoid canal; entotympanic, ectotympanic and pterygoid composing eustachian tube opening (BI); hypoglossal foramen recessed dorsally, lies at same level as jugular foramen; presence of a groove connecting the internal opening of the posterior lacerate foramen to foramen magnum; glenoid fossa ventral to superficies meatus; mediolaterally widened glenoid fossa (BI); smooth posterior surface of glenoid; glenoid fossa wellseparated from porus acusticus (BI); laterally directed root of zygoma; deltoid crest faces anteriorly, reaching the medial side of the humerus in anterior view (BI); strongly marked brachiocephalicus crest of humerus (BI); quadrangular laterodistal corner of scaphoid, in dorsal view (BI); fusion of trapezium and metacarpal I, forming the carpal-metacarpal complex; absence of a strong concavity between greater trochanter and the head of femur (BI); moderately developed lesser trochanter of femur, with a laminar projection; femoral patellar trochlea isolated or only abuts the condylar surfaces; medial trochlear ridge protruded anteriorly to lateral trochlear ridge (BI); distal epiphyses of femur twisted with respect to the main axis; and an obtuse (around 120°) angle formed by facets for cuboid and metatarsal IV, in metatarsal V., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on pages 1526-1527, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Scelidotheriinae PP = 100, age = 17.55 Mya (14.03–20.82). The clade includes the genera Neonematherium, Sibyllotherium, Valgipes, Proscelidodon, Catonyx and Scelidotherium, and was recovered in all analyses performed in this study. The relationships among those genera were consistent among analyses: (Neonematherium, (Sibyllotherium, (Valgipes, (Proscelidodon, (Catonyx, Scelidotherium))))). It was recovered in all but one analysis, par_EW, in which Neonematherium and Sibyllotherium are in a polytomy with the remaining scelidotheriines. Scelidotheriinae was supported by six synapomorphies, all but one recovered by both methods: flat occlusal surface of molariforms; subtriangular Mf1 cross-section (BI); roughly horizontal profile of dorsal surface of the skull, in lateral view; presence of ventral extension in maxilla for dental alveoli; medial palatal process of maxilla extending anterior to lateral process; and well-developed, free-standing paracondylar processes., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1523, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Megatheriidae minus Analcimorphus PP = 100, age = 18.47 Mya (16.97–20.24). This clade is composed of the clades Prepotheriinae, Nothrotheriinae and Megatheriinae, which were recovered in two alternative arrangements: (Prepotheriinae, (Nothrotheriinae, Megatheriinae)) for H models, par_IW10 and par IW5, and (Megatheriinae, (Nothrotheriinae, Prepotheriinae)) for par_EW and par_IW_100. For A models that contain the clade, both arrangements were observed. The clade was supported by 17 synapomorphies (ten for both methods and seven exclusively for BI): Cf1 closer to Mf1 than to anterior edge of maxilla (BI); deep zygomatic process almost covers the entire squamosal exposure; occiput vertical or slightly inclined posterodorsally; sessile occipital condyles (BI); median fusion of posterior alae of vomer; medial expansion of entotympanic dorsal to basicranium; occipital artery passes through a closed canal in mastoid; absence of intertrochanteric ridge; moderately developed lesser trochanter of femur, with a laminar projection (BI); distal epiphyses of femur twisted with respect to the main axis (BI); lateral facet of anterior border of tibial proximal epiphysis located posterior to medial facet (BI); distal fibular articulation for tibia posterolaterally positioned; odontoid process well defined only on distal half of proximodistal length of tibial surface (BI); slightly obtuse (around 120°) angle formed by discoid and odontoid facets of astragalus, in distal view; process for navicular on astragalus at the level of the odontoid facet; transverse diameter of discoid facet of astragalus smaller than that of odontoid facet (BI); and obtuse (around 120°) angle formed by facets for cuboid and metatarsal IV, in metatarsal V.

Lestodontini PP = 100, age = 7.06 Mya (5.32–9.16). The clade is composed of Lestodon and Bolivartherium and was recovered in all analyses. Lestodontini was supported by 11 synapomorphies (six for both methods and five exclusively for BI): presence of a well-developed diastema between Cf1/cf1 and molariforms; Cf1 closer to anterior edge of maxilla than to Mf1; Cf1/cf1 displaced laterally relative to molariform toothrow; anteroposteriorly ovate mf1 cross-section (BI); anteroposteriorly ovate mf2 crosssection (BI); maximum length of nasal bones greater than or equal to three times the width of both nasals (BI); attachment of the base of jugal to skull dorsal to Mf2 (BI); well-developed buccinator fossa of maxilla, with a deep depression; absence of a crest at median suture of palatine; dorsally situated infraorbital canal; and posteroventrally inclined occipital condyle in lateral view (BI).

Nothrotheriinae PP = 100, age = 15.44 Mya (13.02–17.76). The clade is composed of five genera consistently recovered in all analyses with the following arrangement: ((‘ Xyophorus’, Mionothropus), (Pronothrotherium, (Nothrotherium, Nothrotheriops))). Nothrotheriinae was supported by 24 synapomorphies (12 for both methods and 12 exclusively for BI): trapezoidal Mf2 and Mf3 crosssection (BI); absence of a corkscrew-like rotation in the plane of articulation of mandibular condyle, in dorsal view (BI); presence of angle at the junction of symphysis and lower edge of horizontal ramus of mandible in lateral view (BI); large supraoccipital exposure on cranial roof, expanded anteriorly at midline; posterior segments of temporal lines run anterior to but closely parallel the nuchal crest (BI); nasal expands posteriorly, lateral margins divergent (BI); lateral and medial palatal processes of maxilla of equivalent length; interpterygoid and posterior interpalatine regions of roughly equal width; presence of pterygoid/vomer contact (BI); large sinus in pterygoid; alisphenoid-parietal contact present (BI); external occipital protuberance ventral to dorsal nuchal crest, in line with ventral nuchal crest (BI); posterior edge of occipital condyles ends at or anterior to posterior edge of foramen magnum superior border; large exposure of vomer in nasopharynx, covers presphenoid and much of basisphenoid; pterygoid does not participate in bony wall of tympanic cavity; anteroventral process of tegmen tympani as a large bony mass; occipital artery passes through a short canal in mastoid, perforating paroccipital process; glenoid fossa well separated from porus acusticus (BI); anteriorly directed root of zygoma; humeral head widely exposed, raised above the tubercles (BI); proximolateral process of metacarpal II does not extensively overlap metacarpal III proximally, contacting magnum (BI); absence of a detached sulcus delimitating, both medially and laterally, the articular surface of the femoral head in anterior view (BI); absence of a strong concavity between greater trochanter and the head of femur; and femoral patellar trochlea isolated or only abuts the condylar surfaces.

Megatheriinae PP = 100, age = 15.28 Mya (13.30–17.45). The clade, which was recovered in all of our analyses, is composed of the stem genera Diabolotherium and Megathericulus and the clades Thalassocnini and Megatheriini. The arrangement (Diabolotherium, (Megathericulus, (Thalassocnini, Megatheriini))) was recovered in all BI analyses and in par_IW10. In par_EW and par_IW100, the order of divergence between Diabolotherium and Megathericulus is reversed, whereas in par_IW5, Aymaratherium and Thalassocnus are successively sister taxa to Megatheriini, not forming a clade. Megatheriinae was supported by 16 synapomorphies (11 for both methods and five exclusively for BI): Cf1 and cf1 dorsoventrally aligned with molariforms; absence of a well-developed diastema between Cf1/cf1 and molariforms; Cf1 within size range of molariforms; cf1 within size range of molariforms; molariform morphology of Cf1/cf1; trapezoidal Cf1 cross-section; trapezoidal cf1 cross-section; absence of a mandibular fossa posterior to cf1; nuchal crest continuous with dorsal edge of zygomatic process of squamosal (BI); anteroventrally inclined zygomatic process of squamosal; postorbital process roughly at the level of maxillary foramen (BI); distinct neck present at the base of occipital condyles (BI); carotid foramen partially covered ventrally by entotympanic and ectotympanic, in ventral view; glenoid fossa well separated from porus acusticus (BI); absence of entepicondylar foramen of humerus; and medial (trochlear) portion of trochlear notch extends proximal to lateral (capitular) portion (BI)., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1529, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Thinobadistini PP = 81, age = 9.38 Mya (7.52–11.26). The clade is composed of Thinobadistes and Lestobradys and was recovered in all analyses, with the exception of par_EW. Thinobadistini was supported by a single synapomorphy, obtained with both methods: the presence of a diastema between Mf1 and Mf2., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1524, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Choloepodini PP = 98, age = 7.07 Mya (5.67–8.93). This clade is composed of the genus Pliomorphus; the Antillean megalonychines Megalocnus, Parocnus, Neocnus and Acratocnus; and the living sloth genus Choloepus. It was consistently recovered in our analyses with the arrangement (Pliomorphus, ((Megalocnus, Parocnus), (Neocnus, (Acratocnus, Choloepus)))). In one exception—in par_EW— Neocnus, Acratocnus and Choloepus were recovered in a polytomy. Choloepodiniwassupportedbyeightsynapomorphies (six for both methods and two exclusively for BI): snout downturned anteroventrally, in lateral view; evenly convex dorsal surface of the skull; minimum width of palate between toothrows equal or less than width of Mf2; absence of postorbital process of zygomatic arch; occiput wider than deep; presence of recessus meatus (BI); presence of a strong concavity between greater trochanter and the head of femur (BI); and patellar trochlea confluent with lateral, but not with medial condylar surface.

Nematheriinae PP = 100, age = 18.60 Mya (16.85–20.28). This clade contains the genera Nematherium and Analcitherium, and was recovered in all analyses performed in this study. Nematheriinae was supported by seven synapomorphies, all recovered with both methods: anteroposteriorly ovate Mf4 cross-section; snout depressed anteriorly; external nares not greatly enlarged; absence of pterygoid inflation; presence of lacrimal eminence; presence of prominent lateral walls in lacrimal foramen; and jugal and lacrimal anteriorly overlapping facial portion of maxilla., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1523, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Ahytheriini PP = 100, age = 4.44 Mya (1.93–6.98). This clade, composed of the intertropical megalonychids Australonyx, Ahytherium, Megistonyx, Nohochichak and Xibalbaonyx, was consistently recovered in all of our analyses, with two alternative arrangements: (Australonyx, ((Ahytherium, Megistonyx), (Nohochichak, Xibalbaonyx))) in all BI analyses and in par_EW, and (Australonyx, (Xibalbaonyx, (Nohochichak, (Ahytherium, Megistonyx)))) for IW parsimony analyses. Ahytheriini was supported by 23 synapomorphies (13 for both methods and ten exclusively for BI): absence of fossa on palatal surface of maxilla posterior to Cf1; circular Mf1 cross-section (BI); ascending ramus of mandible partially covers posterior teeth in lateral view (BI); coronoid process not hooked posteriorly (BI); posterior external opening of mandibular canal opens anterolaterally, on ascending ramus; high and narrow braincase (BI); nasal uniform width in its posterior half, lateral margins parallel (BI); interpterygoid and posterior interpalatine regions of roughly equal width (BI); large sinus in pterygoid; ventrally situated infraorbital canal; absence of alisphenoidparietal contact (BI); presence of a marked postorbital constriction; postorbital process of frontal roughly at the level of maxillary foramen (BI); triangular occiput in posterior view (BI); occipital condyles situated well dorsal to the dentition (BI); anteromedially orientation of entotympanic (BI); weakly marked brachiocephalicus crest of humerus (BI); humeral head almost hidden behind the tubercles; anconeal process of ulna not extended anteriorly, not overhanging trochlear notch; widest extension of pronator ridge at the proximal half of radial diaphysis (BI); proximal half of femur narrower than distal half; intermediate tibial length, more than two times the width, but less than three times; and fusion of metatarsal I with entocuneiform.

Folivora minus Pseudoglyptodon PP = 100, age = 37.98 Mya (33.43–43.47). The clade is composed of the living genus Bradypus and Eufolivora. It was consistently recovered in all analyses performed in this study, except in par_EW consensus, in which Pseudoglyptodon, Bradypus and Eufolivora were arranged in a polytomy. The clade was supported by nine synapomorphies (two for both methods and seven exclusively for BI): a large basin for the modified orthodentine core; ovate cf1 cross-section (BI); anteroposteriorly ovate Mf1 crosssection (BI); anteroposteriorly ovate mf1 cross-section (B1); bilobate Mf2 and Mf3 cross-section (BI); bilobate mf2 cross-section (BI); ascending ramus of mandible not covering the posterior teeth in lateral view (BI); presence of posterior external opening of mandibular canal; and maximum length of nasal bones greater than, or equal to, twice the width of both nasals, but less than three times (BI)., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1520, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Mylodontini PP = 100, age = 8.19 Mya (6.54–9.95). This clade is composed of ten genera, with the following arrangement in all analyses using H models: (Pleurolestodon, ((Simomylodon, Glossotheridium), (Ocnotherium, (Glossotherium, (Paramylodon, (Oreomylodon, (Mylodonopsis, (Archaeomylodon, Mylodon)))))))). For UN and A models not accounting for ACRV (_e), and MP trees, several alternative arrangements can be observed, regarding the positions of Pleurolestodon, Ocnotherium, Oreomylodon, Glossotherium and Paramylodon, whereas the clades (Simomylodon, Glossotheridium) and (Mylodonopsis, (Archaeomylodon, Mylodon)) are stable. Mylodontini was supported by 11 synapomorphies (seven for both methods and four exclusively for BI): Cf1 smaller than the smallest molariform; cf1 smaller than the smallest molariform (BI); ovate Cf1 crosssection; presence of internal ridge running obliquely or vertically from ventral edge of ascending ramus, near the base of the angle, towards the last tooth; length of coronoid process greater than its height (BI); absence of a mandibular fossa posterior to cf1 (BI); temporal lines laterally situated, do not approximate midline of skull roof; posterior segments of temporal lines run anterior to but closely parallel the nuchal crest; medial palatal processes of maxilla anterior to lateral process; presence of osteoderms; and a slightly obtuse (around 120°) angle formed by discoid and odontoid facets of astragalus, in distal view (BI)., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1525, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236

Urumacotheriinae PP = 60, age = 17.82 Mya (14.03–21.51). This clade was originally proposed with fewer taxa (Negri & Ferigolo, 2004), but was expanded here to include both Urumacotherium and Pseudoprepotherium. It did not receive high node support, despite being present in trees from all analyses. Urumacotheriinae was supported by seven synapomorphies (five for both methods and two exclusively for BI): molariform morphology of Cf1/ cf1 (BI); posterior edge of mandibular condyle inclined posterodorsally; eversion of lateral edge of symphyseal spout (BI); premolariform portion of palate roughly equal or longer than the length of molariform toothrow; presence of a distinct neck at the base of occipital condyles; occipital condyles elongated anteroposteriorly in ventral view; and lesser tubercle of humerus proximally projected as much as greater tubercle., Published as part of Casali, Daniel M, Boscaini, Alberto, Gaudin, Timothy J & Perini, Fernando A, 2022, Reassessing the phylogeny and divergence times of sloths (Mammalia: Pilosa: Folivora), exploring alternative morphological partitioning and dating models, pp. 1505-1551 in Zoological Journal of the Linnean Society 196 (4) on page 1524, DOI: 10.1093/zoolinnean/zlac041, http://zenodo.org/record/7381236, {"references":["Negri F, Ferigolo J. 2004. Urumacotheriinae, nova subfamilia de Mylodontidae (Mammalia, Tardigrada) do Mioceno superior-Plioceno, America do Sul. Revista Brasileira de Paleontologia 7: 281 - 288."]}

In the present paper, we report the occurrence of the cricket subgenus Gryllitara Chopard, 1931 from India along with the description of a new species, Itara (Gryllitara) pilosa sp. nov., from the state of Kerala, South India. Suitable plates with close-up details have been presented to describe the new species, as well as for Itara (Itara) vietnamensis Gorochov being reported from North East India.

Auricularia has a worldwide distribution and is very important due to its edibility and medicinal properties. Morphological examinations and multi-gene phylogenetic analyses of 277 samples from 35 countries in Asia, Europe, North and South America, Africa, and Oceania were carried out. Phylogenetic analyses were based on ITS, nLSU, rpb1, and rpb2 sequences using methods of Maximum Likelihood and Bayesian Inference analyses. According to the morphological and/or molecular characters, 37 Auricularia species were identified. Ten new species, A. camposii and A. novozealandica in the A. cornea complex, A. australiana, A. conferta, A. lateralis, A. pilosa and A. sinodelicata in the A. delicata complex, A. africana, A. srilankensis, and A. submesenterica in the A. mesenterica complex, are described. The two known species A. pusio and A. tremellosa, respectively belonging to the A. mesenterica complex and the A. delicata complex, are redefined, while A. angiospermarum, belonging to the A. auricula-judae complex, is validated. The morphological characters, photos, ecological traits, hosts and geographical distributions of those 37 species are outlined and discussed. Morphological differences and phylogenetic relations of species in five Auricularia morphological complexes (the A. auricula-judae, the A. cornea, the A. delicata, the A. fuscosuccinea and the A. mesenterica complexes) are elaborated. Synopsis data on comparisons of species in the five complexes are provided. An identification key for the accepted 37 species is proposed.

Psectrascelis is one of the most specious genus of darkling beetles of South America, comprising 82 species that inhabit arid and semiarid environments. The species are distributed mainly in Argentina and Chile. In general, the taxonomy of several species in this group has been difficult to approach. An example is Psectrascelis pilosa Solier and Psectrascelis conjugens Fairmaire, species morphologically similar which presents a rather poor original description and mislabelled. These species are distributed sympatrically in coastal terraces at the northern side of Huasco river, northern Chile. We collected several specimens of Psectrascelis on the southern side of Huasco river which are morphologically similar to P. pilosa. Morphological and genetic analysis support the description of the species Psectrascelis ammophila sp. nov., which is more closely related to Psectrascelis elongata and Psectrascelis pilipes specularis than other morphologically similar species (Psectrascelis pilosa and Psectrascelis conjugens). The importance of the Huasco river as barrier and the evolutionary convergence is discussed to explain the crypsis and the phylogenetic pattern found.

The first Oriental species of the genus Trichopsomyia Williston, 1888, Trichopsomyia pilosa sp. nov. (Java), has been discovered and is now described. This Oriental species of Trichopsomyia has several characters strongly differing from the other species within this genus. It is hypothesized that it forms a separate group within Trichopsomyia. One species, Trichopsomyia formiciphila Downes, Skevington & Thompson, 2017, from Australia, is similar to the Oriental species described here, and the group is named after this first described species, hence the formiciphila group. The characters for a future phylogenetic analysis are discussed. The character states of the pilosity of the katepisternum and the shape of the metasternum hitherto used in a phylogenetic analysis of Syrphidae Latreille, 1802 are discussed too.

This study aimed to raise and address nomenclatural issues surrounding Neotropical species in the genus Gnamptogenys, in addition to describing new species accumulated in myrmecological collections. New and reinstated names recognized here include G. pernambucana (Santschi) sp. rev., stat.n., G. lenis sp. n., G. latistriata sp. n., and G. avus sp. n. The queen and intercaste of G. lavra Lattke and the queen of G. pernambucana are described for the first time. For these five species we provide complete descriptions and diagnoses, comments on taxonomy and natural history, distribution data, and high-resolution images, including the first images of G. lavra. An updated identification key for all the Neotropical species of Gnamptogenys is also provided, including the new and revived species as well as G. aspera Morgan and G. pilosa Lattke, not present in previous keys for the genus.

Amblyomma nodosum is a Neotropical tick species distributed from Mexico to Argentina, with adult individuals infesting different mammal species, including anteaters (Pilosa: Myrmecophagidae). Few reports in Colombia have recorded this species in departments such as Meta, Tolima and Valle del Cauca. In this paper we describe this species using taxonomic, morphometric and molecular methods after collecting individuals parasitizing collared anteaters (Tamandua tetradactyla) in the departments of Cundinamarca and Tolima. Adult specimens were identified based on current taxonomic keys and different morphometric variables were documented in nonengorged individuals. Also, DNA was extracted for PCR amplification and sequencing of 16S rDNA, COI and ITS2 genes. Phylogenetic trees were built. One hundred and thirty-four adult ticks were collected and identified as Amblyomma nodosum, based on morphological, morphometric, molecular and phylogenetic analyses. This is the first study to report the presence of this tick species in the department of Cundinamarca, using multiple identification approaches, thus expanding its geographical records in Colombia. Resumo Amblyomma nodosum é uma espécie de carrapato Neotropical distribuída do México à Argentina com indivíduos adultos, infestando diversas espécies de mamíferos, incluindo tamanduás (Pilosa: Myrmecophagidae). Na Colômbia, limitados relatos têm registrado essa espécie em alguns departamentos como Meta, Tolima e Valle del Cauca. Neste trabalho, espécimes foram identificados por meio de métodos taxonômicos, morfométricos e moleculares após serem coletados parasitando indivíduos de tamanduá-mirim (Tamandua tetradactyla) dos departamentos de Cundinamarca e Tolima. Espécimes adultos foram identificados por meio de chaves taxonômicas e diversas variáveis morfométricas foram ristradas em indivíduos não ingurgitados. Além disso, o DNA foi extraído para amplificação pela PCR e por sequenciamento dos genes 16S rDNA, COI e ITS2. Árvores filogenéticas foram construídas. No total, 134 carrapatos adultos foram coletados e identificados como Amblyomma nodosum por meio de análises morfológicas, morfométricas, moleculares e de filogenia. Este é o primeiro estudo que relata, por meio de múltiplas ferramentas de identificação, esta espécie no departamento de Cundinamarca ampliando assim seus registros geográficos na Colômbia.

Nannotrigona Cockerell is one of the few stingless bee genera widely distributed and commonly encountered throughout the Neotropical region. Some species are abundant and managed in meliponiculture. However, the identity and number of species occurring in some countries or areas is almost entirely unknown. Herein we reviewed the species of Nannotrigona in Colombia, a South American country with literature records indicating the presence of either one or five species of this genus. Appraisal of museum specimens revealed the existence of the following eight species, three of which we describe as new: N. camargoi Rasmussen & Gonzalez, N. gaboi n. sp., N. melanocera (Schwarz), N. mellaria (Smith), N. occidentalis n. sp., N. pilosa n. sp., N. schultzei (Friese), and N. tristella (Cockerell).We describe and figure the male of N. camargoi, N. gaboi, N. melanocera, and N. pilosa. In addition, we provide comparative diagnoses, a key to species, and new geographical records.

One new nematode species is described and two new species records are provided from the edge (6,080 m depth) and axis (7,132 m) of Kermadec Trench, Southwest Pacific. Leptolaimus hadalis sp. nov. is characterised by medium body 587–741 μm long, labial region not offset from body contour, inconspicuous labial sensilla, amphid located 12–19 μm from anterior end, female without supplements, male with four tubular precloacal supplements (alveolar supplements absent), tubular supplements almost straight with dentate tip, arcuate spicules and weakly cuticularized dorsal gubernacular apophyses strongly bent distally. In a previously published ecological survey of Kermadec Trench, L. hadalis sp. nov. was the most abundant species in a core obtained at 8,079 m water depth and third most abundant species in a core obtained at 7,132 m, while only one individual was found at 6,096 m depth, and none at 9,175 m depth (Leduc & Rowden, 2018). Alaimella aff. cincta and Desmodora aff. pilosa are recorded for the first time from the Southwest Pacific region. Prior to the present study, Alaimella had only been recorded from coastal locations and from the Weddell sea to a depth of 2,000 m. The record of Desmodora aff. pilosa at 6,080 m depth is the deepest record of a Desmodora species to date, although unidentified Desmodora specimens have been found as deep as 6,300 m in the South Sandwich Trench. The morphology of the Kermadec Trench Alaimella aff. cincta and Desmodora aff. pilosa specimens bear a strong resemblance to their respective type populations from the Northern Hemisphere, but further morphological and molecular data are required to ascertain whether they in fact represent distinct species.

This report continues our monographic analysis of mammalian diversity and Matses ethnomammalogy in the Yavari-Ucayali interfluvial region of northeastern Peru. Based primarily on specimens collected in the region from 1926 to 2003, interviews with Matses hunters, and published sight surveys of large mammals, we document the local occurrence of 33 species of xenarthrans, carnivores, perissodactyls, artiodactyls (including cetaceans), and sirenians. All of the species in these groups, with the exception of the Amazonian manatee (Trichechus inunguis), are recognized and named by the Matses, from whom we recorded extensive accounts of mammalian natural history.The local xenarthran fauna consists of nine species (Cabassous unicinctus, Priodontes maximus, Dasypus novemcinctus, D. pastasae, Bradypus variegatus, Choloepus hoffmanni, Cyclopes didactylus, Myrmecophaga tridactyla, Tamandua tetradactyla), all of which are represented by examined specimens. Only two xenarthrans (D. pastasae and C. hoffmanni) are primary game species for the Matses, who are familiar with many aspects of their biology that were previously unrecorded in the scientific literature. However, Matses interviews also provide important new information about the behavior of D. novemcinctus (a secondary game species) and M. tridactyla, neither of which has previously been studied in rainforested environments.The local carnivore fauna consists of 16 species (Atelocynus microtis, Speothos venaticus, Leopardus pardalis, L. wiedii, Panthera onca, Puma concolor, Pu. yagouaroundi, Eira barbara, Galictis vittata, Mustela africana, Lontra longicaudis, Pteronura brasiliensis, Bassaricyon alleni, Nasua nasua, Potos flavus, Procyon cancrivorus), most of which are represented by examined specimens; six species without preserved voucher material are known from camera-trap photographs and/or unambiguous sightings by Matses hunters and field biologists. Although the coati (N. nasua) is the only carnivore occasionally hunted by the Matses for food, Matses interviews are richly informative about the natural history of other species, notably including S. venaticus, Leopardus spp., Pa. onca, Puma spp., and E. barbara.All of the local ungulates (Tapirus terrestris, Pecari tajacu, Tayassu pecari, Mazama americana, M. nemorivaga) are hunted by the Matses for food, and the hunters we interviewed are correspondingly well informed about the natural history of most of these species, with the exception of the seldom-encountered gray brocket (M. nemorivaga). Both species of local cetaceans (Inia geoffroyi, Sotalia fluviatilis) are familiar to the Matses, although neither is eaten.The xenarthrans, carnivores, ungulates, and aquatic mammals that inhabit the Yavari-Ucayali interfluve are all widespread species, so this component of the regional fauna, as currently understood, is not biogeographically distinctive, nor is it extraordinarily diverse (by western Amazonian standards). Although we discuss several noteworthy taxonomic and nomenclatural issues relevant to these taxa, the principal contribution of this report consists in the natural history information compiled from our Matses informants and the resulting overview of local community structure as defined by diurnal activity, locomotion, social behavior, and trophic relationships.

The complex of Uapaca kirkiana is revised in the northwest of the Zambezian region (Angola, D. R. Congo, Zambia) based on herbarium taxonomy. Five taxa are recognised, i.e. Uapaca kirkiana Müll. Arg. var. kirkiana, U. kirkiana var. gossweileri (Hutch.) Meerts, U. kirkiana var. benguelensis (Müll. Arg.) Meerts, U. teusczii Pax f. teusczii and U. teusczii f. pilosa (P. A.Duvign.) Meerts. A determination key is provided. Specimens lacking curly hairs on leaves have long been misinterpreted or overlooked. The name Uapaca teusczii Pax is resuscitated to accommodate such specimens. New diagnostic characters of U. teusczii are highlighted including a glabrous ovary, anthers with ciliate connectives and the pubescent calyx of female flowers. U. teusczii is a characteristic species of the wet miombo, widespread in Katanga and Kwango (D. R. Congo), extending locally to Angola and N Zambia. A lectotype is selected for U. albida, U. homblei, U. kirkiana var. benguelensis, U. kirkiana var. gossweileri, and U. teusczii f. pilosa; second-step lectotypification is provided for U. munamensis., info:eu-repo/semantics/published

Ptilothyris Walsingham, 1897, a poorly known Afrotropical genus of Lecithoceridae, is revised. Eight new species are described and illustrated: P. subcucullata Park, sp. nov., P. crassiella Park, sp. nov., P. drepanodes Park, sp. nov., P. enormisella Park, sp. nov., P. hylodes Park, sp. nov., P. leifaarviki Park, sp. nov., P. pilosa Park, sp. nov., and P. vokaensis Park, sp. nov. Each species is compared with type specimens of the related species deposited in various museums worldwide. COI barcode sequences of six species were successfully obtained and are provided. Of the 13 previously known African species of the genus, P. serangota ab. cyanea Ghesquière, 1940 is synonymized with P. serangota Meyrick, 1932, and the following three species are excluded from the genus: brachysema Meyrick, 1938, crocophracta Meyrick, 1938, and nemophorella Ghesquière, 1940. The latter three will be transferred to related genera in forthcoming studies. Adults and genitalia of all the known species including the new species are illustrated, and a key to the species is provided.

Se actualiza la información taxonómica y geográfica de Ochthera para Colombia, un género con especies benéficas, reconocidas por sus hábitos depredadores de insectos pequeños terrestres y larvas de mosquitos. Se incluye el primer registro de Ochthera (O.) pilosa y nuevos datos de distribución para Ochthera (O.) regalis y Ochthera (O.) angustifacies. Se agregan ilustraciones, comentarios de su distribución y mapas. We update taxonomic and geographic information for genus Ochtherain Colombia, a taxon containing benefi cial species known by their predatory habits on terrestrial insects and mosquito larvae. We consider a fi rst record of Ochthera (O.) pilosa, and new distribution data of Ochthera (O.) regalis and Ochthera (O.) angustifacies, adding illustrations, comments on their distribution, and maps. Incluye referencias bibliográficas

This study describes and illustratesChrysosplenium macrospermumY.I.Kim & Y.D.Kim, a new plant species from Changbaishan Mt. (Baekdusan Mt.) in northeastern China. The species is most similar toChrysosplenium valdepilosumin the seriesPilosabut is readily distinguishable by short arching sterile branches, multiple (up to 3) flowering stems, and smooth surfaced seeds (without tubercles), which are ca. 30–50% larger than those of other members in the series.

Males of the neotropical bee genus Ruizantheda Moure sensu lato (Halictinae: Caenohalictini) differ from those of other caenohalictine genera in having the outer gonostylar plate with a large membranous region that extends to the ventral region, and the ventral gonostylar lobe retrorse. These features permit the placement of six additional species described here, four of which are new, in the genus in this broader sense. The males of these six species share the following characteristics: profemur swollen; mesofemur exceedingly swollen; mesotibia slightly swollen with flat, minutely ridged ventral area and small apical tooth; S4 shortened medially, commonly largely hidden under S3, with transverse median depression and apical margin emarginate; apicolateral lobes of S4 with tuft of branched setae and stout, simple, recurved setae on margin; S5 with U-shaped gradulus and setal patch, sometimes inconspicuous; outer lateral expansion of penis valve bilobed, and median posterior margin of volsella strongly angulate. Together, these species constitute a new group within Ruizantheda and share with the previously known Ruizantheda inca Coelho, Felizardo, & Engel, R. aerugineus Coelho, Felizardo, & Engel, and R. kallos Coelho, Felizardo, & Engel, a short outer gonostylar plate, terminating before the apex of the main gonostylar lobe as well as the presence of long setae on its dorsal surface. In other species of Ruizantheda the outer gonostylar plate extends beyond the apex of the main gonostylar lobe and lacks setae. The species comprising this distinctive subgroup of Ruizantheda, and treated herein, are: R. nigra n. sp., R. colombiana n. sp., R. venezuelana n. sp., R. baeri (Vachal) n. comb., R. pilosa n. sp., and R. gaullei (Vachal) n. comb. Two of the new species expand the distribution of the genus northward into Colombia and Venezuela. In addition to the description and illustration of the species, an updated key to the species of Ruizantheda s. l. is provided.

The cockroach genus Eupolyphaga is revised. Twenty species and two subspecies are recognized, with thirteen species and two subspecies are new to science: E. daweishana sp. nov., E. dongi sp. nov., E. fengi sp. nov., E. hanae sp. nov., E. hupingensis sp. nov., E. maculata sp. nov., E. nigrifera sp. nov., E. nigrinotum sp. nov., E. pilosa sp. nov., E. robusta sp. nov., E. shennongensis sp. nov., E. wooi sp. nov., E. xuorum sp. nov., E. everestiana reni subsp. nov., and E. fengi yongshengensis subsp. nov. Homœogamia sinensis Saussure is placed as a junior synonym of E. sinensis (Walker) and the replaced name E. limbata (Kirby) for Homœogamia sinensis is invalid, the status of Eupolyphaga thibetana (Chopard) is recovered. Male genitalia of species in the genus is described and illustrated. Females and oothecae of some species are described and illustrated. Distribution maps and a checklist of Eupolyphaga are provided. A key to males of Eupolyphaga is given. Plenty habitat photographs are shown.

Myrmecophaga tridactyla Linnaeus, 1758 Figure 4 Records. First record was in August 08, 2016, camera trap 18 (25°03.37ʹ S, 049°59.44ʹ W) and subsequently in the same monitoring site. Identification. Two species of anteaters are known in the region of PNCG, with each having different body size and color pattern of pelage. Myrmecophaga tridactyla is a large species and weighs up to 45 kg. It has small ears and small eyes, long snout and an extremely long tongue; the thick pelage varies from dark gray and black, but with the paws white paws and some black bands at the top. The tail is robust and covered by hair., Published as part of Pereira, Alan Deivid, Bazilio, Sergio & Orsi, Mário Luís, 2018, Checklist of medium-sized to large mammals of Campos Gerais National Park, Paraná, Brazil, pp. 785-799 in Check List 14 (5) on page 790, DOI: 10.15560/14.5.785

Two new species of Polana DeLong, 1942 are described and illustrated, P. (Angusana) pilosa sp. nov. from Peru and P. (Declivella) maculosa sp. nov. from Brazil. Polana (Striapona) desela Delong, 1979 is redescribed; the female terminalia of this species are described and illustrated for the first time.

5. Red Silky Anteater Cyclopes rufus French: Myrmidon roux / German: Rotlicher Zwergameisenbar / Spanish: Oso hormiguero sedoso rojo Taxonomy. Cyclopes rufus Miranda et al., 2017, “Porto Velho, Rondonia, Brazil (08°51°167S; 064°00’53"W).” This species is monotypic. Distribution. W Amazon Basin, in interfluve between Rio Madeira and Rio Aripuana; N limit is possibly Amazon River, and S limit is unknown. Descriptive notes. No specific data are available. Color of the Red Silky Anteater is unique; dorsum has a distinct reddish tone, and tail and limbs are more yellowish red. Ventral and dorsalstripes are absent. Fronto-nasal region of skull is not depressed, giving a straight profile. External aperture of ear is directed laterally. Naso-maxillary sutures are about parallel, with wide fronto-maxillary suture. Fronto-parietal suture is trapezoidal, and pterygoid bone does not overlap tympanic bulla. Habitat. Amazonian forest. Food and Feeding. The Red Silky Anteater is an opportunistic forager. Its diet consists almost entirely of ants. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The Red Silky Anteater is found in areas between Madeira and Aripuana rivers that suffer from environmental degradation and effects of large hydroelectric dams. The Red Silky Anteater is probably threatened with extinction, and its distribution and population status, considering recent taxonomic changes, need to be urgently evaluated. Bibliography. Hayssen et al. (2012), Miranda et al. (2017)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Cyclopedidae, pp. 99-102 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 101, DOI: 10.5281/zenodo.6627824

4. Thomas's Silky Anteater Cyclopes thomasi French: Myrmidon de Thomas / German: Thomas-Zwergameisenbér / Spanish: Oso hormiguero sedoso de Thomas Taxonomy. Cyclopes thomas: Miranda et al., 2017, “Porto Walter, Acre, Brazil.” Specific name honors O. Thomas in recognition of his extensive contribution to mammalogy and specifically taxonomy of Cyclopes. Monotypic. Distribution. W Amazon Basin, from Rio Jurua SW to Rio Ucayali region, in Pasco and Ucayali provinces (EC Peru); E limits are unknown but might not extend beyond Rio Madeira. Descriptive notes. No specific data are available. Body color of Thomas's Silky Anteater is strikingly orange to reddish brown; legs and tail are gray. Dorsal stripe is absent. Ventral stripe is little developed and faint. Fronto-nasal region of skull is not depressed, giving a straight profile. External aperture of ear is directed anteriorly. Naso-maxillary sutures diverge proximally, with very short fronto-maxillary suture. Fronto-parietal suture is triangular, and pterygoid bone does not overlap tympanic bulla. Habitat. Amazonian forests. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The [UCN Red List. It is necessary to evaluate conservation status of Thomas’s Silky Anteater because of recent taxonomic revisions, requiring clarification ofits distribution. Bibliography. Miranda et al. (2017)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Cyclopedidae, pp. 99-102 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 101, DOI: 10.5281/zenodo.6627824

Russell A. Mittermeier, Don E. Wilson (2018): Bradypodidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 118-132, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6621602

Russell A. Mittermeier, Don E. Wilson (2018): Myrmecophagidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 88-90, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6656622

Russell A. Mittermeier, Don E. Wilson (2018): Myrmecophagidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 88-90, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6656622

Russell A. Mittermeier, Don E. Wilson (2018): Cyclopedidae. In: Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos. Barcelona: Lynx Edicions: 99-102, ISBN: 978-84-16728-08-4, DOI: http://doi.org/10.5281/zenodo.6627824

3. Pale-throated Three-toed Sloth Bradypus tridactylus French: Paresseux a trois doigts / German: \WeiRkehl-Faultier / Spanish: Perezoso tridactilo de garganta péalida Other common names: Pale-throated Sloth Taxonomy. Bradypus tridactylus Linnaeus, 1758, “America meridionalis arboribus.” Restricted by O. Thomas in 1911 to Suriname. This species is monotypic. Distribution. N South America, from E Venezuela (E & S of the Orinoco River), E through the Guianas to N Brazil (W to the Amazon River Delta and bounded in the S by the Amazon River). Descriptive notes. Head-body 450-755 mm, tail 22-110 mm, ear 5-21 mm, hindfoot 40-120 mm; weight 3.4-6.5 kg. Adult males are smaller than females. The Pale-throated Three-toed Sloth has long, coarse, dark gray hair with black spots, especially around back and shoulders. Pelage is greenish because of algae that grows along cracks of hair shafts. Throat and head are yellowish. Males have dorsal speculum with orange-yellow patch and broad black central stripe. Males and females have small vestigial tails. Females have two mammae on their chests. Chromosomal complement is 2n = 52 and FN = 56. Habitat. Tropical forests with diverse tree species and complex horizontal and vertical cover. Although seldom seen, the Pale-throated Three-toed Sloth is abundant throughout its distribution. Food and Feeding. The Pale-throated Three-toed Sloth is a strict folivore, consuming leaves from the canopy of overstory trees. Its diet has been reported to almost exclusively contain leaves of Cecropia spp. (Urticaceae), butit also eats leaves of Ceiba samauma (Malvaceae), Elizabetha paraensis (Fabaceae), and Hevea viridis (Euphorbiaceae). The Pale-throated Three-toed Sloth has one of the slowest rates of digestion and fermentation because of the large volume of digesta it retains in its gut and its low metabolic rate. Breeding. Breeding of the Pale-throated Three-toed Sloths is seasonal. Pregnancy, inferred from timing of births and assuming six-month gestation, occurs in August— February. One very small young is born per litter, and births appear to occur during the long rainy season (March—June). Interbirth interval is c.12 months, and age of independence for juveniles is ¢.5 months. Young cling to venters of their mothers. They sample leaves as early as three weeks old but do not consume substantial amounts of leaves until they are c.5 weeks old. Young are closely associated with their mothers for the first year oflife. Activity patterns. Pale-throated Three-toed Sloths are strictly arboreal and do notlive outside of the forest. Similar to their congeners, they have limited activity; one individual was observed resting/inactive for 18-5 hours/day, hanging upside down from branches. While sleeping, they are most often seated in crooks of trees on horizontal branches, with their heads tucked into their chests under their arms. They have been observed sunning themselves in tree canopies. The Pale-throated Three-toed Sloth swims well and can use rivers as dispersal corridors. Movements, Home range and Social organization. Pale-throated Three-toed Sloths are primarily solitary, although groups of up to three individuals have been observed. Individuals can be aggressive—primarily with their claws—to conspecifics if they occupy the same tree. They can reach high densities within Neotropical forests. In Suriname, it was the most abundant non-volant mammal and constituted a large proportion of total mammalian biomass in the forests. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Palethroated Three-toed Sloth has a relatively wide distribution, occurs in protected areas, and is rarely targeted for consumption or human use. Clearing of Neotropical forests is detrimental to Pale-throated Three-toed Sloths because they are vulnerable to habitat loss and fragmentation. Bibliography. Beebe (1926), Eisenberg & Thorington (1973), Foley et al. (1995), Hayssen (2009b), Richard-Hansen & Taube (1997), Taube, Keravec et al. (2001), Taube, Vié et al. (1999), Thomas (1911b)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Bradypodidae, pp. 118-132 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 131, DOI: 10.5281/zenodo.6621602

2. Pygmy Three-toed Sloth Bradypus pygmaeus French: Paresseux nain / German: Zwergfaultier / Spanish: Perezoso tridactilo pigmeo Other common names: Pygmy Sloth Taxonomy. Bradypus pygmaeus R. P. An- derson & Handley, 2001, “Panama: Bocas del Toro: Isla Escudo de Veraguas, West Point.” Taxonomic status unclear; it was described as a species based on morphometric analyses, notably a smaller body size, but results of recent molecular studies are conflicting and could be a pygmy form of B. variegatus. Monotypic. Distribution. Restricted to the Escudo de Veraguas I, 18 km offshore from the Valiente Peninsula, on the Caribbean coast of Panama. Descriptive notes. Head-body 485-530 mm, tail 45-60 mm, ear 10 mm, hindfoot 94-110 mm; weight 2:5.3-5 kg. The Pygmy Three-toed Sloth is similar in color to the Brown-throated Three-toed Sloth (B. variegatus); it is gray with long, coarse hair, covered with algae causing hair to appear green. The Pygmy Three-toed Sloth has dark orbital stripe over lighter colored and browner face. Males have dorsal speculum of shorter cream to orange hair. Forelimbs and hindlimbs have three large and distinct claws. Habitat. [sland mangrove forests. Pygmy Three-toed Sloths are believed to be primarily associated with red mangroves (Rhizophora mangle, Rhizophoraceae) that occur along the margins of Isla Escudo de Veraguas. A few sloths have been observed in interior, upland forests of the island. More systematic efforts are needed to survey interior forests to determine the degree to which Pygmy Three-toed Sloths are associated with mangrove and other tree species. Food and Feeding. The Pygmy Three-toed Sloth has been observed eating leaves from mangroves and other species (e.g. Cecropia spp., Urticaceae). Breeding. No information. Activity patterns. Pygmy Three-toed Sloths are largely arboreal, but terrestrial and even swiming locomotion has been recorded. Movements, Home range and Social organization. One study of Pygmy Three-toed Sloths systematically searched all mangrove stands (10-7 ha in total on the island) and located 79 individuals for a density of 7-4 ind/ha. The island is 430 ha, butit is unknown to what extent upland forests are used by Pygmy Three-toed Sloths. A study recently compared sleep characteristics between Pygmy Three-toed Sloths and Brown-throated Three-toed Sloths and found that while both species slept for the same amount of time (9-10 hours/day), the Brown-throated Three-toed Sloth preferred to sleep at night; Pygmy Three-toed Sloths showed no preference. It was suggested that the difference could be the result of different predation risk, with island Pygmy Three-toed Sloths being exposed to less risk than mainland Brown-throated Three-toed Sloths. Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. Given the extremely limited distribution of the Pygmy Three-toed Sloth on a single, small island and uncertainty around the overall population size, exact distribution, and threats of habitat loss from visitors to the island, the viability of the population is unclear. Bibliography. Anderson & Handley (2001), Hayssen (2008), Kaviar et al. (2012), Voirin (2015), Voirin et al. (2014)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Bradypodidae, pp. 118-132 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 131, DOI: 10.5281/zenodo.6621602

6. Xingu Silky Anteater Cyclopes xinguensis French: Myrmidon du Xingu / German: Xingu-Zwergameisenbar / Spanish: Oso hormiguero sedoso de Xingu Taxonomy. Cyclopes xinguensis Miranda et al., 2017, “Vitoria do Xingu, Para, Brazil (Usina Belo Monte) (03°17°12”S; 051°53’48"W).” This species is monotypic. Distribution. E Brazilian Amazon, limited in N by Amazon River, in W by Rio Madeira, and in E by Rio Xingu; S limit is unknown. Descriptive notes. No specific data are available. Dorsal color of the Xingu Silky Anteater is gray, rump is yellow, venter is pale yellowish, and legs and tail are gray. Dorsalstripe is evident, and ventral stripe is indistinct and irregular. Fronto-nasal region of skull is not depressed, giving a straight profile. External aperture of ear is directed laterally. Naso-maxillary sutures are about parallel to each other, with wide fronto-maxillary suture. Fronto-parietalis triangular, and pterygoid bone overlaps tympanic bulla. Habitat. Amazonian forests. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The Xingu Silky Anteater lives in an area with substantial environmental degradation and is probably threatened with extinction. Its distribution and population status, considering recent taxonomic changes, need to be urgently evaluated. Bibliography. Miranda et al. (2017).

1. Brown-throated Three-toed Sloth Bradypus variegatus French: Paresseux a gorge brune / German: Dreifinger-Faultier / Spanish: Perezoso tridactilo de garganta marrén Other common names: Bolivian Three-toed Sloth, Brown-throated Sloth Taxonomy. Bradypus variegatus Schinz, 1825, “Stidamerika.” Restricted by R. Mertens in 1925 to “Brasilien (wahr- scheinlich Bahia)” (= Brazil, probably Bahia). There have been up to nine subspecies recognized, but given their distribution, geographical variability, cryptic nature, and lack of clearly differentiated forms, the actual number is unclear, and subspecific taxonomyrequires reassessment. Distribution. From Honduras SE throughout Central America to Colombia and W & S Venezuela, and S into Ecuador, Peru, most of Brazil, N Bolivia, Paraguay, and N Argentina. Descriptive notes. Head-body 519-540 mm, tail 52-55 mm, ear 8-22 mm, hindfoot 85-180 mm; weight 3.7-6 kg. Adult Brown-throated Three-toed Sloths weigh less than they appear due to long and thick pelage. Males are slightly smaller (less than 10%) than females. They are gray, with long, coarse hair, but hair is often covered with mutualistic algae (and other microbiota), causing it to have green tinge. The Brown-throated Three-toed Sloth has a dark orbital stripe over a lighter colored and browner face, giving the appearance of a mask. Males have dorsal speculum of shorter cream to orange colored hair, with dark stripe running vertically down the center of it. Contrasting dorsalstripe appears to become accentuated with age and is sometimes accompanied by black spots on either side. Males and females have small vestigial tails. Forelimbs and hindlimbs have three large and distinctive claws. The Brown-throated Three-toed Sloth has long forelimbs (370-450 mm) and relatively short hindlimbs (320-370 mm). Chromosomal complement is 2n = 54-55 and FN = 56-58. Habitat. Tropical forests with diversity oftree species and complex horizontal andvertical cover. The Brown-throated Three-toed Sloth can reach high densities in agricultural landscapes if overhead canopy trees are maintained. It is a generalist and uses manydifferent tree species, although individuals will specialize on a few species of trees and even spend a disproportionate amount of time resting and foraging within a few individual trees, or “modal” trees, in their home ranges. In an agricultural landscape in Costa Rica, Brown-throated Three-toed Sloths were strongly dependent on intact tropical forest and two species oftrees in the nettle family (Cecropra obtusifolia and Coussapoa villosa, Urticaceae). In general, trees that have crowns exposed to sunlight and those that contain masses oflianas are selected. Food and Feeding. The Brown-throated Three-toed Sloth is a strict folivore, consuming leaves in canopies oftrees. Individuals typically consume leaves in trees where they spend a considerable amount oftime. Diets vary spatially and overall include leaves from more than 50 plant species. They prefer and mostly consume young leaves, presumably because of their nutritional value. The Brown-throated Three-toed Sloth has one of the slowest rates ofdigestion. Nevertheless, estimated leaf removal rate is 5-1 g ofleaves/kg/day. Sloths, in general, are foregut-fermenting mammals, and the Brownthroated Three-toed Sloth possess a unique gut community dominated bythe bacterial phyla Proteobacteria and Firmicutes. Three-toed sloths possess a highly conserved, low-diversity foregut community, with highly abundant species of Neisseria bacteria. Breeding. The Brown-throated Three-toed Sloth is polygynous, with males excluding male competitors from their core home ranges. In one study in Costa Rica, only 25% of resident adult males sired offspring, and one individual male sired 50% of all juveniles. Sedentary lifestyle of the Brown-throated Three-toed Sloth appears to facilitate polygyny because multiple females can persist within small patches of habitat and be monopolized by a single male. Females appear to use important mating strategies by switching mates among breeding seasons and shifting their home ranges during estrus. The Brown-throated Three-toed Sloth breeds seasonally. Gestation is ¢.6 months in length, and females give birth to one young over a six-month period that peaks in late February. Neonates are dependent on their mothers, clinging to their venters for 100 days or more. This relatively short time to offspring independence allows females to breed in most years; on average, nearly 75%ofadult females produce young in the wild, and this frequency appears to vary little from year-to-year. Juvenile survival can be high but is lowest immediately following maternal independence. Activity patterns. Brown-throated Three-toed Sloths are active diurnally and nocturnally, but they are more often active during daylight hours and generally avoid activity during the crepuscular periods of the day. Adults usually sleep for 15-18 hours/day, either holding on to a tree with their claws or resting in the crotch of a tree with their heads tucked into their chests. Three-toed sloths have a high degree of heterothermy in their body temperature, and therefore, they behaviorally adjust to ambient temperatures throughout the day. Individuals can be observed in the mornings ascending to the top of the canopy, presumably to warm in full sunlight, and descending into shade as daytime temperatures increase. During rainstorms, Brown-throated Threetoed Sloths do not seek shelter but typically rest in place. Movements, Home range and Social organization. Brown-throated Three-toed Sloths climb slow and deliberately and position themselves on undersides of branches with their bodies perpendicular to branches when moving. In a single day, an individual only moves an average of 40 m and is often found on the same tree. Once a week, individuals descend to the forest floor. While on the ground, they create shallow depressions with their vestigial tails and defecate. This ritualized behavior reinforces sloth moth (Pyralidae) colonization of fur and increases nitrogen and phosphorous levels in sloth fur contributing to algal growth. Home range sizes do not differ between sexes or by breeding status and are 0-1-19 ha. Most adult male Brown-throated Three-toed Sloths maintain exclusive use of core areas in their home ranges. Home ranges of males overlap with many females but not each other, and home ranges of adult females overlap each other. Juveniles strongly selectfor forest cover during dispersal, avoiding agricultural habitat types. Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Brown-throated Three-toed Sloth has the widest distribution of the four species of Bradypus and occurs across various forest types. It occurs in protected areas and is rarely targeted for consumption or other human uses, although occasionally it is hunted in Central America and South America by indigenous groups and for the pet trade. Clearing of tropical forests for crops grown in monoculturesis particularly detrimental because these plantations retain little forest structure. Brown-throated Threetoed Sloths are more vulnerable to habitat loss and fragmentation than other sloths, especially two-toed sloths (Choloepus spp.). Nevertheless, previous work has shown that shade-grown agriculture, like that found in certain types of cacao plantations embedded within an agroecosystem, can support viable populations of the Brown-throated Three-toed Sloth, if immigration from surrounding forests is maintained. In particular, intact riparian forests provide critical movement corridors for maintaining connectivity of populations in modified tropical landscapes. Bibliography. Dill-McFarland et al. (2016), Garcés-Restrepo et al. (2017), Hayssen (2010), Mendoza etal. (2015), Mertens (1925), Montgomery & Sunquist (1978), Moraes-Barros etal. (2010), Noss et al. (2008), Pauli & Peery (2012), Pauli, Mendoza et al. (2014), Pauli, Peery et al. (2016), Peery & Pauli (2014), Sunquist & Montgomery (1973b), Svartman (2012)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Bradypodidae, pp. 118-132 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on pages 130-131, DOI: 10.5281/zenodo.6621602

Family CYCLOPEDIDAE (SILKY ANTEATERS) • Smallest extant anteaters with silvery gray to golden brown to reddish bodies and limbs and darker dorsal, sternal, or ventral streaks and lighter extremities in some species; myrmecophagous; and arboreal. • 20-40 cm.• Nearctic (marginally) and Neotropical Regions. Rainforest to mangrove forests. • 1 genus, 7 species, 7 taxa. • Nospecies threatened; none Extinct since 1600., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Cyclopedidae, pp. 99-102 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 99, DOI: 10.5281/zenodo.6627824

2. Linnaeus’s Two-toed Sloth Choloepus didactylus French: Paresseux a deux doigts / German: Eigentliches Zweifingerfaultier / Spanish: Perezoso didactilo de Linneo Other common names: Linné’s Two-toed Sloth, Southern Two-toed Sloth Taxonomy. Bradypus didactylus Linnaeus, 1758, “in Zeylona.” Corrected by O. Thomas in 1911 to Suriname. Subspecific taxonomy requires reassessment; different authors indicate potential differentiation between western and eastern populations. Monotypic. Distribution. NE & S Venezuela (Orinoco Delta and S of the Orinoco River), the Guianas, N & NC Brazil (along the Amazon River W to Maranhao and S to Para and Mato Grosso states), and into upper Amazon Basin of SW Colombia (N up to Meta and Guainia departments), E Ecuador, and NE & E Peru;its S limit in W Amazonian Brazil is unclear. Additional visual records are described for SE Para State and need to be confirmed. Descriptive notes. Head—body 540-880 mm, tail 9-33 mm, ear 20-38 mm, hindfoot 103-170 mm; weight 4-11 kg. There is no outward sexual dimorphism. Linnaeus’s Two-toed Sloth is the largest extant sloth species. It has long foreand hindlimbs, nearly equal in length. Forelegs have two large, curved claws, and hindlimbs have three. Teeth are rootless and ever-growing and lack enamel. Incisors are absent, and premolars are indistinguishable from molars. Pelage ranges from tan to buffy brown and, in the wild, can appear shaded pale green from algae that grows on hair. Face is light brown,is not furred, and has leathery texture. Throat hair is similar in color to pectoral hair, and ventral hairis lighter than dorsal hair. Underfuris lacking, and abdominal hair is parted in middle and flows toward mid-dorsum. Young Linnaeus’s Two-toed Sloths tend to have soft and darker pelage. Diploid numbers range between 2n = 52-54 and 2n = 64-65. Habitat. Primary and secondary tropical, moist lowland forests but up to elevations of 2400 m. Food and Feeding. Linnaeus’s Two-toed Sloth is an arboreal herbivore and generalist. Its feeding behavior is poorly known, and most information comes from captivity. In captivity, they consume an average of 350 g/day of solid food. Coprophagy was observed in captivity. Linnaeus’s Two-toed Sloths were observed feeding at human latrines in north-eastern Peru, which could supply needed nutrients such as salt or protein from human feces. Breeding. Linnaeus’s Two-toed Sloths reproduce year-round. Estruslasts 10-14 days, and ovarian cycles based on hormones are 4-5 weeks. Gestation lasts 330-350 days, and interbirth intervals are 14-27 months. Females give birth to single young and are not sexually mature before c¢.3 years old; males do not breed before 4-5 years old. Mating occurs abdomen to abdomen and takes place either on the ground or while hanging from a branch. Linnaeus’s Two-toed Sloths live up to 27 yearsin captivity, and one female gave birth at 25 years old. Mortality of infants in captivity is high. Females carry their infants on their ventral or dorsal regions; infants become independent at c.5 months old. Activity patterns. Linnaeus’s Two-toed Sloths are nocturnal. They spend most of their time in forest canopiesto forage,rest, sleep, and breed. They descend to the ground to defecate every 3-5 days. In captivity, they spend 45% oftheir time sleeping, 25% foraging or grooming (mostly self-grooming), 20% resting with eyes closed, and 10% awake but inactive. When resting, they assume a sitting position hanging onto overhead support with one or more limbs or supported by hindlimbs while head and forelimbs are curled ventrally over groin. Movements, Home range and Social organization. Linnaeus’s Two-toed Sloths are mostly solitary. They move between trees through interconnecting tree crowns or using lianas. Density in French Guiana is 0-5 ind/ha. In captivity, Linnaeus’s Two-toed Sloths seem to tolerate being kept in pairs or small groups; however, fighting between two males, probably for access to a female, was observed in captivity. Status and Conservation. Classified as Least Concern on The IUCN Red List. Overall population trends of Linnaeus’s Two-toed Sloths are still unknown, but they are unlikely to be declining fast enough to represent a conservation threat. Linnaeus’s Twotoed Sloths occur in many protected areas, but they are probably hunted opportunistically. Considering variation in karyotypes, a taxonomic review is needed to assure appropriate conservation action. Bibliography. Adam (1999), Chiarello & Please (2014), Eisenberg & Redford (1999), Gardner & Naples (2008), Gilmore et al. (2000, 2001), Hautier et al. (2016), Heymann et al. (2011), Jones (1977), Moraes-Barros & Arteaga (2015), Rezende et al. (2013), Taube et al. (1999), Thomas (1911b), Trinca et al. (2006), Troll, Gottschalk, Heuer et al. (2015), Troll, Gottschalk, Seeburger et al. (2013), Voirin et al. (2013), Wetzel (1985a)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Megalonychidae, pp. 104-117 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 117, DOI: 10.5281/zenodo.6647064

2. Northern Tamandua Tamandua mexicana French: Tamandua du Mexique / German: Nordlicher Tamandua / Spanish: Oso melero septentrional Other common names: Mexican Anteater Taxonomy. Myrmecophaga tamandua var. mexicana Saussure, 1860, “Tabasco,” Mex- 1CO. The name 7. mexicana was first used by R. M. Wetzel in 1975. In 1982, Wetzel recog- nized five subspecies, but A. L. Gardner in 2005 accepted only four. Both species of Tamandua are mostly allopatric, but in some areas of the extreme northern South America, they may occur in sympatry. Four subspecies recognized. Subspecies and Distribution. T.m.mexicanaSaussure,1860—SMexico(fromColimaandMichoacanstatesinthePacificslopeandSanLuisPotosiStateintheGulfside)andthroughoutCentralAmericaStoHonduras. T:m.instabilisJ.A.Allen,1904—NColombia(N&EofEAndesRange)andNWVenezuela. T.m.opistholeucaGray,1873—SHondurastoWColombiaandWEcuador;alsoonBarroColoradoI(Panama). T: m. punensisJ. A. Allen, 1916 — W & SW Ecuador (including Puna I) and NW Peru. Descriptive notes. Head-body ¢.560 mm, tail 400-675 mm, ear 40-46 mm, hindfoot 80-110 mm; weight 3-6 kg. The Northern Tamandua is not outwardly sexually dimorphic. It is medium-sized but slightly smaller than the Southern Tamandua (7: tetradactyla). Head of the Northern Tamandua is slender and intermediate in size between those of the Giant Anteater (Myrmecophaga tridactyla) and the Common Silky Anteater (Cyclopes didactylus). Rostrum is more than 40% oftotal length of cranium. Skull of the Northern Tamandua has four pairs of orbital foramina, but there are usually three in Southern Tamandua. Ears of the Northern Tamandua are smaller than those of the Southern Tamandua. Eyes and mouth are tiny, the latter without teeth. The 400mm tongue is contractile and has circumvallate papillae that produce sticky saliva. Northern Tamanduas have distinctive black vests on paler fur background (tan or yellowish). Dense short hair covers body. Muscles of forelimbs are strong and well adapted to tear apart nests of prey. There are four claws on forefeet, with second and third well developed. Northern Tamanduas walk on sides of their hands, with toes inward and distributing weight on thickened tissue of the pad. Hindfeet are plantigrade and have five smaller claws. Prehensile tail is nearly naked for the last three-quarters ofits length, and it has 40-42 caudal bones, more than in the Southern Tamandua (31-39). Chromosomal complementis 2n = 54, FN = 104. Habitat. Wide variety of habitats from open savannas, thorn scrublands, and mangroves to forests (lowland, gallery, evergreen, and deciduous tropical) and humanaltered areas. Northern Tamanduas use terrestrial habitats and forest canopies, where they spent almost one-half of their time when active. Food and Feeding. Northern Tamanduas exploit terrestrial and arboreal nests of ants and termites. Prey is found with a keen sense of smell; prominent claw is used with a twisting motion to break the nest. Northern Tamanduas can eat ¢.9000 insects/ day. One larvae and pupae are consumed for every nine mature ants; immature forms have higher caloric values than adults, so they are nutritionally important. Northern Tamanduas exploit 50-80 nests of ants or termites daily. They eat ant genera Camponotus, Crematogaster, Azteca, Procryptocerus, and Dolichoderus and termite genera Nasulilermes and Microcerotermes. Northern Tamanduas also eat palm fruits (Attalea butyracea, Arecaceae) in Panama. Breeding. Northern Tamanduasare solitary but come together to breed. A wild mating pair was observed in Costa Rica. When the pair met, the male sniffed female's rump and then followed her; on several occasions, he touched her from behind. Sometimes, the female was antagonistic to the male, including standing up bipedally in defense. Two copulations were observed in five minutes, and each lasted 10-30 seconds. Mating is aseasonal. Gestation lasts 130-150 days. Northern Tamanduas commonly give birth to one young that is cared for by its mother for c.1 year. Young may be left for short periods while the mother forages. Longevity is 9-5 years. Activity patterns. Northern Tamanduas are scansorial, and they can exploit arboreal and terrestrial habitats. They can be active for up to eight hours, day, or night. When they are active, 40% of the time is spent in trees, and 90% ofrest sites are in trees, tree hollows, and liana tangles; they also rest in abandoned armadillo burrows. Northern Tamanduas maintain ¢.20 different restsites in core areas of their home ranges, and they use 1-2 of them each day. Features of restsites include high tree densities (especially palms), heights, and dense vegetation. Trees are preferred as restsites probably to avoid predation from Ocelots (Leopardus pardalis), Pumas (Puma concolor), and Jaguars (Panthera onca). Movements, Home range and Social organization. Northern Tamanduas moved arboreally from one tree crown to another, spending about three minutes in each tree and visiting ¢.20 trees/hour. Lianas and thick branches facilitate progress. When Northern Tamanduas are active, movement is continuous, except for rests of ¢.30 minutes; long feeding bouts are related to finding high caloric prey. Average daily movement is 1000 m, but up to 3400 m have been recorded. Northern Tamanduas swim skillfully, with snouts held above the water. They are most active in peripheries of home ranges. Home range size was 0-25 km? in Panama, and overlap can occur near edges. In Costa Rica (Punta Leona Private Refuge) and Panama (Barro Colorado Island), densities were very low at 0-0006 ind/km? and 0-0005 ind/km?, respectively. In coastal forest of Cabo Blanco, Costa Rica, Northern Tamanduas are described as abundant. Status and Conservation. CITES Appendix III (Guatemala). Classified as Least Concern on The IUCN Red List. The Northern Tamandua is considered common across most of its distribution, but in the northern part in Mexico, it is endangered. Risks to some populations include deforestation, habitat loss, fires, poaching, road kills, and conflict with dogs. In some regions of Mexico, traditional people eat Northern Tamanduas, and Yucatan people use their claws in cock fighting. They are illegally kept as pets. More research is needed for a better conservation assessment of the Northern Tamandua. Bibliography. Aguiar & Fonseca (2008), Alfredo & Cuaron (1987), Allen (1904, 1916), Brown (2011), Brown, LaPoint et al. (2012), Brown, Montgomery et al. (2014), Burton & Ceballos (2006), Ceballos & Arroyo-Cabrales (2012), Esser et al. (2010), Gardner (2005, 2008), Genoways & Timm (2003), Jorge & Pereira (2008), Lubin (1983), Lubin & Montgomery (1981), Matlaga (2006), Montgomery (1985b, 1985c), Montiel et al. (2000), Navarrete & Ortega (2011), Ortega et al. (2014), Redford & Dorea (1984), Rodrigues et al. (2008), Rossoni et al. (1981), SEMARNAT (2010), Silveira (1968), Superina et al. (2010), Timm et al. (2009), Timock & Vaughan (2002), Tlapaya & Gallina (2010), Urquiza-Haas et al. (2011), Wetzel (1975, 1982, 198b5a)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Myrmecophagidae, pp. 88-90 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 89, DOI: 10.5281/zenodo.6656622