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Toromys albiventris, sp. nov. Echimys rhipidurus: Sanborn, 1949: 286 (a misidentification). Makalata sp. nov. Emmons, 2005: 279. HOLOTYPE: FMNH 55483, an adult male skin and skull, collected by José Schunke, 15 August 1946. The tail of the holotype is slightly bent at midpoint. TYPE LOCALITY: Peru: Ucayali, Yarinacocha (8.35° S. 74.59° W; 155 m). ETYMOLOGY: From the Latin albus (“white”), and venter (“belly”). The only Toromys with white underparts. DIAGNOSIS: The smallest Toromys, with greatest length of skull 51.6–54.2 mm, and maxillary toothrow 10.1–11.9 mm (tables 5, 6). Dorsum coarse brown agouti, finely lined with black, tinted faintly rusty from behind shoulder to rump; aristiform hairs composed of narrow bristles with whiplike tips, mixed with some wider, spinelike aristifoms, also with hairlike tips (fig. 4F). Underparts sharply demarcated white, yellow-white, or beige-white, with faintly buffy areas (fig. 16). Tail shorter than head and body length; tail hair distal to the extension of the dorsal hair onto the base longer basally but does not cover scales, diminishes in length and pales in color down the length of the tail, tip thinly haired, with short, fine brown hairs. Incisors white or pale yellow. Nasal bones flare laterally posteriorly, to a variable extent. Mandible slender, with a shallow sigmoid notch, and coronoid process small and low. DESCRIPTION: Dorsum coarsely grizzled brown-agouti, lined with black; back posterior to shoulders, flanks, ventral tail base, and hind limbs, are tinged rusty reddish; forequarters and head tinged more ochraceous yellow lined with black; extreme tip of muzzle dull rusty in some specimens, including the holotype (fig. 16), but barely so in others (fig. 13A). Dorsal hairs a mixture of several widths of wider and narrower bristlelike aristiforms, pale gray basally, banded black or dark brown, then buff or rusty, with dusky, buff, or rusty tips that are drawn out into long hairlike processes (fig. 4F). Hind feet rusty buff above, toes whitish; forefeet gray above. Claws unpigmented and small for the genus, silvery ungual tufts reach beyond their tips. Dorsal hair extends about 3.5 cm onto the tail base, colored like the back above, dull, unbanded rusty below. Posterior to the extension of body hair onto the tail base, tail hair is monocolor black to brown dorsally, beige to white ventrally, over pigmented skin. Hairs longest at the dorsal base and midline, producing an indistinct keel to halfway down the tail, black or brown for about basal one-third, paling and diminishing gradually in length. Length of the tail hair varies between syntopic individuals, from distinctly to barely elongated. The longest hairs near the tail base reach across about five scale rows. Most tail hairs emerge from the scales in pairs, one longer and robust, the other shorter and finer. Tail scales squarish to hexagonal, about as long as wide, in uneven rows on dried skins. Underparts, including the sparsely haired inner limbs, white or slightly tinged yellowish or buff, with a grayish to gray-brown lateral band at the junction with the dorsal pelage. Throat and lower neck crossed by a faint buffy collar. One specimen, LSUMZ 8904, has ventral hairs with dusky subterminal bands lending a brownish tinge to the abdomen and with pale brownish bands across the lower throat and abdomen between the flanks, but the inner limbs are monocolored dirty white. Ear pinnae small, thinly haired inside with buffy hairs, with whitish or pale gray hairs behind; a few inconspicuous salient hairs extend from the rims. Mystacial vibrissae robust and black, the longest reach to the shoulder when laid back; genal and supercilliary vibrissae fine and inconspicuous; antebrachial vibrissae present. A female has two pairs of used lateral mammae, one above the flank and the other about midway between fore- and hind limbs and about 1.5 cm into the dorsal pelage field. The dried hindfeet appear unpigmented and relatively narrower across the base of the digits than are those of T. rhipidurus; they have small tubercles around and between the plantar pads, and collars around the digital pads, which appear large, but details of the feet are unclear. Forefeet with abundant small tubercles on the midline of the palm. Cranium in dorsal view with a long, slender braincase and a short rostrum; nasal bones pinched medially and flared laterally posteriorly and slightly anteriorly (fig. 9C). Parietal-interparietal suture nearly straight or bowed posteriorly; interparietal wide dorsally across the whole width of the posterior cranium. Incisive foramina slightly ovoid, widest anteriorly, with the maxillary and premaxillary portions of the septum broadly fused, and with a strong medial ridge leading onto the palate to between the premolar roots (figs. 7C, 17C). Supraorbital ridges beaded posteriorly, moderately expanded into shelves. Jugals dorsoventrally narrow, with jugal fossa narrow, and a slender insertion onto the maxillary. Incisors are white to pale yellow. Occlusal pattern as for the genus, but lophs and flexi of more uneven widths and angles, and M1 and M2 mures nearly at center of teeth. Unworn upper cheek teeth divided in half by the mesoflexus into two joined pairs of lophs (fig. 12A, B). COMPARISONS WITH OTHER SPECIES: Toromys albiventris differs from T. rhipidurus by having more coarsely grizzled upperparts with more rust-tinged dorsum behind the shoulders (fig. 16), less rusty muzzles, shorter tail hairs, a white or pale-beige venter, slender foreclaws, and narrower hind feet. Compared to T. rhipidurus the cranium is more slender, with a narrower rostrum, narrower mesopterygoid notch, and the medial parietal-interparietal suture curved posteriorly rather than straight or recurved anteriorly. LOCALITIES AND NATURAL HISTORY: The known specimens are from two localities (fig. 1). The type locality, Yarinacocha, is a left-bank oxbow lake of the Río Ucayali, a widely meandering, whitewater river, at about 150 m. Some of the Yarinacocha specimens were collected by J.M. Schunke in 1946, and the others were collected in the 1960s by expeditions from Louisiana State University. The second locality, “Suaya,” is the origin of two specimens collected in 1926 by J.M. Schunke. Vaurie (1972) spelled this locality as “Suayo” and provided approximate coordinates of 07°18′ S; 74°55′ W, which is downstream and east of Contamana on the right bank of the Río Ucayali. We here emend this locality to Lagos del Suhaya, 7.46° S; 75.18° W; 160 m, left bank of the Río Ucayali, about 20 km WSW of Contamana. These are a group of lakes in a landscape depression that borders the meander zone of the river, and are likely accessible by boat during high water. This locality is on current maps, and is plausible because Schunke collected 10 days later at “Pampa Hermosa,” a currently occupied and mapped village about 35 km downriver (north) of Contamana. Moreover, it is pronounced the same way as the original was written on the specimen label. None of the specimens are accompanied by habitat notes, but A.L. Gardner, collector of LSUMZ 14409 and LSUMZ 14408 (a Makalata), recalled in person to L.H.E. that both specimens were shot at night from the forest canopy, along the raised bank bordering a narrow channel leading into the west side of Yarinacocha lake, on the northern outskirts of Pucallpa. The forest was disturbed, dense secondary regrowth, with largediameter lianas. The two species were taken in the same stretch of trail. The region around Pucallpa is among the most heavily populated and deforested of the Peruvian Amazon, and now is crisscrossed by roads, plantations, and reservoir dams, and scarred from recent burns (Google Earth, 2017). Some gallery forests remain, likely because they are flood prone, and T. albiventris could be present in these. However, the right (east) bank of the Ucayali remains nearly entirely forested. Yarinacocha is a well-known ornithological locality from many LSUMZ expeditions (Stephens and Traylor, 1983). REMARKS: Sanborn (1949) identified the 1946 FMNH specimens as Toromys (“ Echimys ”) rhipidurus, but noted them as enigmatic, remarking correctly that the tail hair differed among specimens of the series. The two 1926 AMNH specimens are browner, with red-brown tail hairs paler than on those collected in 1946, and we suspect the older specimens have bleached to more orange hues. One collected in 1968 (LSUMZ 14409) has black dorsal tail hairs, while the 1962 specimen (LSUMZ 8904) has red-brown hairs that are scarcely elongated, but that form a slight dorsal keel along the tail. LSUMZ 8904, taken 31 July 1962, was multiparous and pregnant with a single, 30 mm embryo. SPECIMENS EXAMINED: PERU: Ucayali, Yarinacocha (FMNH 55483, 55484, 55485, 55486, 55487, 62100, 62099, LSUMZ 8904, LSUMZ 14409*); Lagos del Suhaya (AMNH 98666*, 98667)., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 39-42, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Sanborn, C. C. 1949. Mammals from the Rio Ucayali, Peru. Journal of Mammalogy 30: 277 - 288.","Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press.","Vaurie, C. 1972. An ornithological gazetteer of Peru (based on information compiled by J. T. Zimmer). American Museum Novitates 2491: 1 - 36.","Stephens, L., and M. A. J. Traylor. 1983. Ornithological gazetteer of Peru. Cambridge, MA: Museum of Comparative Zoology (Harvard University)."]}

Pattonomys punctatus (Thomas, 1899) Loncheres punctatus Thomas 1899: 153 (original description). TYPE SPECIMEN AND TYPE LOCALITY: The holotype (by original designation, BMNH 98.12.1.18), an adult male skin and skull, one of three specimens collected 2 July 1898 by G.K. and S.M. Cherrie, at “Caicara, Orinoco,” Bolívar, Venezuela. 4 In the 1830s de Blainville chaired Natural History and Anatomy at MNHN, and I. Geoffroy St.-Hilaire noted that he owed de Blainville and Roulin for “communication” of the specimens to him (1838: 887). He later added that those specimens were poorly preserved and half putrified, but that afterward he received another in good condition (presumably after describing the species) from which he redescribed it in much greater detail (1840: 422). Cécile Callou (MNHN) has kindly researched the history and provenence of MNHN-ZM-MO-1995-1397, which when examined in 1999 by L.H.E. was labeled A-7276 (an 1850 catalog number) and two other numbers. It has now been recataloged as a syntype, with a new number (C. Callou, personal commun.). DIAGNOSIS AND DESCRIPTION: The largest species of the genus in nearly all measurements (table s 5, 6). Head grizzled gray, a mixture of black finely streaked with white, with large whitish postauricular patches. Pelage of back pale gray-based, yellowish agouti streaked with black, with rusty tints on posterior middorsum, rear of rump slightly darker than midback. Blackbanded, white-tipped aristiforms on rump only (two specimens from Apure), or a few also on midback (the holotype and a specimen from Bolívar); fine underhairs rusty. Sides of head and shoulders, and tops of feet, grizzled gray. Underparts sharply demarcated pale gray or white with pale buff on anterior and inguinal areas; inner limbs white. Tail clothed distally with fine dusky hairs, slightly paler below. Cranium robust, with wide, strongly developed supraorbital shelves angled upward and raised above the frontals in old individuals; parietal crest strongly developed. Zygomatic arches strongly bowed outward in dorsal view (fig. 9E). Jugal robust and dorsoventrally deep. Upper incisors orthodont (fig. 7E). Masticatory and buccinator foramina separate (fig. 15A). Palatal notch wider than in other congeners, with outwardly bowed sides bordering a wide canal. Molar rows slightly diverge posteriorly to accommodate the wider mesopterygoid fossa (fig. 10E). Palate wide, about as wide as the molars between the roots of M1. Incisive foramina narrow and parallel sided. Auditory tympanic bullae not inflated, the lateral profile of the anterior bulla nearly flat (fig. 7E). GEOGRAPHIC RANGE AND HABITAT: Known only from Venezuela near the Rio Orinoco and its tributaries in Apure and Bolívar states. Perhaps from gallery forests, as the localities are near rivers in regions generally of seasonally flooded grassland (llanos). SPECIMENS EXAMINED: VENEZUELA: Bolívar, Caicara, Río Orinoco (BMNH 98.12.1.18 [holotype]), 95.12.1.20, 52.12.3.21, 93.12.1.19; AMNH 171389, 171391, 171392, 29872 *, 29873); Hato La Florida, 35 km ESE Caicara (USNM 406402); Apure, 38 km NNW Puerto Paez, Río Cinaruco (USNM 374741, 374742 *)., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 30-31, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Thomas, O. 1899. On new small mammals from South America. Annals and Magazine of Natural History (7) 3: 152 - 155."]}

Pattonomys flavidus (Hollister, 1914) Loncheres flavidus Hollister 1914: 143 (original description). TYPE SPECIMEN AND TYPE LOCALITY: The holotype (by original designation, USNM 63218) consists of the skin and skull of an adult male with shot damage to the head, collected by W. Robinson on 9 July 1895 at “ El Valle, Margarita Island,” Nueva Esparta, Venezuela. DIAGNOSIS: Perhaps as large as Pattonomys punctatus, cranium intermediate in size between P. punctatus and P. carrikeri (based on one individual, tables 5, 6). Dorsal pelage grizzled ochraceous yellowish lined with black, slightly browner over the rump. White-tipped spines robust and confined to the rump. Underhairs ochraceous yellow. Head and neck grayish; underparts yellow-buff. The postauricular patches seem gray-white, but head skin badly damaged. Feet gray above. Cranium with zygomatic arches tapering anteriorly, not bowed outward; masticatory and buccinator foramina separate. Palatal notch V-shaped, with straight borders; bullae not inflated (9F, 10F). Mandible robust and deep below the toothrow (fig. 7F: arrow). REMARKS: Based on the holotype, Pattonomys flavidus is not more clearly aligned in morphology with any one of the other three species. We failed to obtain good mitochondrial sequences for P. flavidus, however, some 100 bp Illumina reads indicate a close relationship to its geographical neighbor P. carrikeri (sequences identical), but with insufficient data to place it definitively, so we provisionally maintain it as an independent lineage. This island endemic is apparently represented in collections only by the 1895 type specimen. Three recent specimens were karyotyped, but not preserved (Aguilera et al., 1998). Nonetheless, they prove that the species was extant in the mid-1990s. GEOGRAPHIC RANGE AND HABITAT: The type locality is El Valle, on the eastern end of Margarita Island, Venezuela, but recent specimens were from the Macanao Peninsula at the western end of that island (Aguilera et al., 1998). The lowlands of the island are largely deforested, but there is a forested protected area on the western side, where the recent specimens were captured, and the highlands of the eastern side, the provenance of the holotype, remain patchily forested (Google Earth, 2017). This continental-shelf island is separated from the mainland by two shallow (Nueva Esparta, Margarita Island, El Valle (USNM 63218 * [holotype])., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 32-33, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Hollister, N. 1914. Descriptions of four new mammals from tropical America. Proceedings of the Biological Society of Washington 27: 141 - 144.","Aguilera, M., N. Sangines, and A. Perez-Zapata. 1998. Echimys semivillosus, a rodent species with a very high chromosomal number. Caryologia 51: 181 - 187.","Guilderson, T., L. Burckle, S. Hemming, and W. Peltier. 2000. Late Pleistocene sea level variations derived from the Argentine shelf. Geochemistry, Geophysics, Geosystems 1: Paper number 2000 GC 000098."]}

Pattonomys Emmons, 2005 TYPE SPECIES: Nelomys semivillosus I. Geoffroy St. -Hilaire, 1838. CONTENTS: Four valid species, as diagnosed below. DIAGNOSIS: A genus of large, arboreally adapted Echimyidae (tables 5, 6). Dorsal pelage with strong, wide, gray-based aristiform spines that narrow to sharp tips; some spines entirely black or dusky tipped, others with dark subterminal bands and contrasting white tips that impart a pronounced speckled appearance (fig. 4B). Prominent pale postauricular patches (fig. 14B). Cranium generally like that of Makalata and Echimys, short and broad, with more or less expanded, winglike supraorbital shelves that curve upward from the frontal bones (figs. 7E–H, 9E–H, 10E–H; see also Emmons, 2005: figs. 11, 14). Upper cheekteeth with widely open flexi, greatly anteroposteriorly expanded protocone and posteroloph (probably metaloph and posteroloph combined), and a markedly oblique paraflexus on dP4 (fig. 12E–G). These expanded lophs are accentuated with wear. The four lophs of dP4–M2 are markedly unequal in length and width: the anteroloph is broad and squared lingually, tapering labially; and the posteroloph is pointed labially and strongly curved along the posterior margin of the tooth. Lower premolar tetralophodont, with two labial and two lingual flexids (fig. 12E–G). DESCRIPTION: Spines wide, sharp, and stiff, with tips not drawn out into long hairlike processes but end in microscopic hairs (fig. 4B). White-tipped spines are sprinkled singly among the lateral and dorsal pelage from shoulder or midback to rump, increasing posteriorly in number and in the length of pale tips. Pale-tipped spines sparsely and evenly distributed over the speckled rump. Dorsal spines mixed with fine, ochraceous yellow to orange setiform bristles and sparse dull reddish or yellow-ochraceous underhairs; pelage slate-gray based. Head, neck, and shoulders grizzled gray lined with black, in a mixture of long, narrow, dusky and whitish aristiforms (fig. 14B). Sides grayish from head to flank between gradually or sharply demarcated pale venter and yellowish or reddish-brown tinged middorsal pelage. Ear tips with an inconspicuous, sparse fringe of long fine, salient hairs. Dorsal pelage extends onto the tail base to about the hind feet of study skins; distal tail evenly covered above and below to its tip with fine, monocolored white to red-brown hairs that do not cover scales but curl outward, so that the tail looks distinctly hairy (fig. 5A). Vibrissae robust, dense, and black, the longest reaching to shoulder when laid back. Two pairs of lateral mammae lie in the dorsal pelage field, one about midway between the limbs, the other just anterior to the flank. Hind feet, based only on P. carrikeri, with four small, ovoid digital pads and a fifth small outer pad budding laterally from D4. Hypothenar pad joined to large naked heel pad with only a slight depression between them (fig. 6B); thenar pad elongate and slender, separated from the joined hypothenar and heel pads by a deep, narrow groove containing palmar tubercles. Plantar surfaces between pads with a few inconspicuous tubercles; digital pads bordered by distinct rings or collars of tubercles (fig. 6B); forefoot digital pads also bordered by indistinct collars of tubercles, but the central palm with few or none. On the cranium, the frontal-squamosal suture at the back of the orbit is raised in a sharp ridge (fig. 7E–H). Mastoid processes short, to the middle or lower edge of the auditory meatus, adpressed to the bulla, and enclosed to the tip by occipital bone (fig. 7E–H) and see also Emmons (2005: 263, fig. 5). Postglenoid foramen enclosed below by the petrosal bone in an anteriorly facing tube; the salient ventral lip of the auditory meatus is usually visible in dorsal aspect (fig. 9G), and is further extended in mature individuals by two overlapping additional bony rings that cup the meatus ventrally and laterally, but are incomplete dorsally (fig. 15A). These are not fused and can be lost in skin preparation. Nasal bones in dorsal aspect flared distally, slightly pinched in medially, and about squared or slightly rounded posteriorly (fig. 9E–H). Incisive foramina small, narrow, and slightly ovoid, with deep grooves and ridges leading onto the anterior palate to between the first or second cheekteeth (fig. 10E–H). Maxillary toothrows nearly parallel. Mandible with a low coronoid process only slightly shorter than the narrow condyloid process, and a shallow sigmoid notch (fig. 7E–H). Mandibular foramen in a fossa arising low on the condyloid ridge; masseteric crest not curved upward anteriorly, but forms a nearly straight line from the base of the angular process (fig. 7E–H). Incisor enamel yellowish white, lowers tinged more orange than uppers; upper incisors opisthodont, robust, and strongly curved, with the base of the roots outside of, or level with, the maxillary base of the zygoma. Maxillary cheekteeth (fig. 12E–G) squarish to rectangular, with rounded posterior lophs and an uneven appearance due to unequal loph sizes. DP4–M3 always with two labial and two lingual flexi. Anterior lingual flexi (paraflexi) short, reaching only about half of the width of the tooth; the posterior lingual flexi (hypoflexi) longer; both labial flexi (para + hypoflexi) extend slightly more than half of the width of the tooth and are of the same length. DP4 and M1 always with a mure in the center of the tooth; M2–3 are with or without a mure, and all four flexi slant posteriorly from the edge to the middle of the tooth, so that labial and lingual flexi slant slightly in opposite directions. If without a mure, the joined flexi have a sharp elbow where the mure would be (fig. 12G, arrow). Lower premolar usually divided by one central flexid into two V-shaped lophs, but with wear forms a midtooth mure. In m1–2 the lingual flexids are parallel and of about equal length. Flexids of all mandibular cheekteeth wide. Hypoconid with a flat labial profile (fig. 12E–G). Posterior borders of lower cheekteeth straight. Phallus of P. carrikeri long and slender, with a long, pointed, bacular papilla, small urethral lappet, border of ventral crater wall a straight line without a U- or V-shaped ventral fold (Emmons, 2005: fig. 10C); those of other species are undescribed. Karyotypes of four specimens, which from their localities we assign to P. flavidus and P. carrikeri, were identical, 2 N = 94, among the highest found in Mammalia (Aguilera et al., 1998). COMPARISONS WITH OTHER GENERA: A white-speckled, spiny pelage is unique to the genus, although Leiuromys, Mesomys, Lonchothrix, and Phyllomys pattoni have strong, buff-tipped spines. There is no geographic overlap of these genera with Pattonomys. The maxillary occlusal pattern of Pattonomys species is unique for the family, and even single or worn upper teeth usually can be identified to genus. A quadralophodont lower premolar is shared only with Leiuromys among Echimyini, but in that genus, the mure is near the lingual edge of the upper molars, and the posterior edges of the lower cheekteeth are curved, while the mure of Pattonomys species is near the center of the upper molars, and the posterior edges of the lower molars are straight. The flexids of Pattonomys are wider than those of other Echimyini. Likewise, the long, thin, simple phallus of P. carrikeri is thus far unique among the Echimyini; those known for the other genera are short, thick, and with additional folds (Leite, 2003; Emmons et al., 2005: fig. 10C), but descriptions are needed for other species in every genus., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 27-29, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press.","Aguilera, M., N. Sangines, and A. Perez-Zapata. 1998. Echimys semivillosus, a rodent species with a very high chromosomal number. Caryologia 51: 181 - 187.","Leite, Y. L. R. 2003. Evolution and systematics of the Atlantic tree rats, genus Phyllomys (Rodentia, Echimyidae), with description of two new species. University of California Publications in Zoology 132: 1 - 118 + xv."]}

Toromys rhipidurus (Thomas, 1928) Echimys rhipidurus Thomas, 1928: 291 (original description). TYPE SPECIMEN AND TYPE LOCALITY: The holotype (by original designation, BMNH 28.7.21.89), an adult male skin and skull collected by R. W. Hendee on 25 January 1928 at “ Pebas. Up. Amazons, 300′,” Loreto, Peru. DIAGNOSIS AND DESCRIPTION: Much smaller than Toromys grandis (tables 5, 6), dorsum finely grizzled yellow-brown agouti heavily lined with black, sides paler yellowish; rostrum in front of and below the eyes, inner rear of thighs, dorsal and ventral tail base, and sometimes a lateral line are dull rust red-orange or red tinged (figs. 13B, 14C, 16). Small, pale yellow postauricular patches are prominent in life, but not evident on most specimen skins (fig. 14C, 16). Aristiforms narrow, black or banded with ochraceous-buff subterminal bands, tipped dusky or ochraceous yellow (fig. 4E). Underparts pale gray-brown, sometimes with a white band between the elbows or white chest patches (fig. 16); gray color imparted by white-based and pale-tipped hairs with wide subterminal gray bands. Tail distal to the dorsal extension of body hair onto the base with long, monocolor black or dark rust-brown hairs above, which can nearly cover the scales proximally, but which shorten and thin distally to expose the scales at the tail tip, such that the tail appears conical; tail hairs paler laterally; self rusty at base below. Scales at dorsal tail base longer anterior-posteriorly than wide laterally, with strongly curved posterior edges (like fish scales, fig. 5C). Feet brown to grayish above; with sparse ungual tufts shorter than the strong claws. Plantar surfaces black pigmented, as for genus described above. Eyes mahogany brown, with pointed-ovoid, vertically slit pupils. Cranium robust, rostrum short and broad, and nasals slightly pinched in medially (fig. 9B). Jugal fossa deep and pointed anteriorly, with a strong ventral lip. Postglenoid fossa with or without a strong beaded lip of petrosal bone below. The space between the auditory meatus and ridge of squamosal above it as wide as or nearly the width of the meatus. Squamosal usually extends as a triangular tongue into the top of the mastoid process (fig. 7B). Medial parietal-interparietal suture between the temporal ridges generally strongly recurved anteriorly or with a medial notch (12/17), but can be straight (5/17, including the holotype). Upper molars with narrow lophs at right angles to the cranial axis, lophs notably parallel and of even width, and traversed by narrow parallel flexi in a distinctly laminar occlusal pattern (fig. 12D). First two lower molars with straight, angled posterolophids (fig. 12D). Incisor faces deep to medium orange; the upper incisor roots arise posterior to the anterior base of the maxillary zygoma. NATURAL HISTORY. Little has been recorded of the habits of this species, other than that it is generally arboreal and can use tree hollows. A number of localities where specimens were collected are in white sand, “ varillal ” habitat (Río Nanay), a specialized ecosystem that harbors endemic plants and vertebrates (e.g., Alonso and Whitney, 2001). SPECIMENS EXAMINED: PERU: Loreto, Pebas (BMNH 28.7.21.89 [holotype]); Río Napo, Lago Mirario (not located), (BMNH 32.8.4.21, 32.8.4.22); Boca Río Mazán, (BMNH 32.8.4.22a); Maynas, Puerto Indiana; (AMNH 73231, 73266–73277 *, 73791); Orosa, Amazon River, (AMNH 74084–74086 *); Genaro Herrera (= Jenaro Herrera), (AMNH 276710); Iquitos, Pampa Chica, (FMNH 87243, 87244 *, 87245–87248); Río Nanay, Santa Rita, (FMNH 87249); Río Manití, Santa Cecilia (FMNH 87249–87252); Maynas, Quistococha, (FMNH 122991, 122992); Nazareth, Río Yavary, (FMNH 19854)., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 38-39, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Thomas, O. 1928. The Godman-Thomas expedition to Peru. - viii. On mammals obtained by Mr. Hendee at Pebas and Iquitos, upper Amazons. Annals and Magazine of Natural History (10) 2: 285 - 294.","Alonso, J. A., and B. M. Whitney. 2001. A new Zimmerius tyrannulet (Aves: Tyrannidae) from white sand forests of northern Amazonian Peru. Wilson Bulletin 113 (1): 1 - 9."]}

Leiuromys, new genus TYPE SPECIES: Echimys occasius Thomas, 1921: 450. CONTENTS: Only the type species. ETYMOLOGY: From the Greek leios (“smooth, bald”) and uros (“tail”), thus, “bald-tailed mouse,” in reference to the slick, hairless appearance of the tail, which is more naked than that of any other arboreal echimyine except lowland species of Dactylomys. DIAGNOSIS: Small, arboreally adapted brown rats, heavily spined dorsally with abundant wide, sharp, spines on midback from neck to rump, spines intermixed with fine wavy rust-red underhairs (fig. 3; tables 3–6). Tail hairs short, fine, and inconspicuous, so that the tail looks bare and even shiny. Auditory tympanic bullae small, flattened in lateral aspect. Inferior jugal process reduced to nearly absent, but jugal fossa deep and sharply pointed anteriorly. Masticatory and buccinator foramina joined in a single aperture. Incisors bone white; upper incisor root does not extend to within the maxillary base of the zygoma. Lower premolars tetralophodont. Third upper molars trilophodont. The hypoflexids on m1–m3 fall nearly opposite the middle of the mesolophids, such that the teeth viewed from the lingual side are nearly symmetrical, rounded M-shapes. Upper lingual and lower labial hypoflexi short and wide, about a quarter of the width of tooth, but labial meso- and metaflexi long, such that when worn to a mure in M1 and M2, the mure is on the far lingual edge of the tooth. All mandibular cheekteeth have curved posterior borders. DESCRIPTION: Upperparts with abundant, wide, strong spines on midback from neck to rump, spines intermixed with fine, wavy, rust-red underhairs that show through aristiforms, giving back a rusty hue (fig. 3). Spine tips sharp, narrowing abruptly, with short microscopic hairlike processes at tips (fig. 4A). Spines tipped with pale buff sparsely speckle the sides of the hindquarters and tail base. Dorsal spines pale gray-brown at base, with a dusky band, a narrow, subterminal pale whitish or buffy band, and dusky, buff, or black-brown tips (fig. 4A). Upperparts uniform light ochraceous brown, between Cinnamon and Sayal Brown (Smithe, 1975), lined with dusky, or blackish Dusky Brown (the holotype and AMNH 71897). The spines on the middorsum from neck to rump are the darkest and nearly unbanded (figs. 3, 4A). Head and muzzle brown above, without any red color. Cheeks below the ears, forelimbs, and hindlimbs tinged grayish. Feet pale beige to rusty above; silvery ungual tufts reach to the ends of the claws. Ventral pelage of soft aristiforms, colored strong to pale buff, grading gradually from the sides (fig. 3); inner thighs slightly darker pinkish buff; the chest and lower abdomen and inguinal regions can have self-white patches, including the tail base, or are crossed by a white band between the elbows. Vibrissae black, moderately robust and the longest reaches to the shoulder. Ears short, pigmented, and nearly naked, lined with a few wispy, dusky hairs. Tail slightly longer than head and body length, brown pigmented, with a naked and somewhat shiny appearance. Dorsal hair ends abruptly close to the body at the tail base; distal tail hairs inconspicuous and closely adpressed. Scales on the tail base evenly hexagonal, wider than long, tapered at the lateral ends. Dorsally each scale has one robust, scalelike dark hair about two and a half scale rows long, and one shorter, fine, hair (fig. 5B). Ventrally, tail hairs somewhat longer (three scale rows), adpressed, and silvery. An adult female (FMNH 84259) has two used lateral pairs of mammae spaced about evenly between the limbs and 1.5 cm into the dorsal pelage field, and possibly an unused inguinal pair. As far as can be seen on dried specimens, the feet have small digital pads collared with rings of tubercles as in Toromys species (fig. 6A), and with numerous plantar tubercles on the bottom and sides of the feet around the pads. Cranium lightly built for the subfamily, inferior jugal process much reduced or essentially absent (figs. 7D, 8, 9D, 10D). Mastoid processes are short, reach to the middle of the auditory meatus, and are adpressed to the cranium; the space above the auditory meatus is nearly as wide as the meatus. Parietal ridges not raised posterior to the postglenoid fossa. Masticator and buccinator foramina joined in a narrow, horizontal slit or a large oval foramen (MCZ 37964) (fig. 8B). Posterior opening of the alisphenoid canal thin walled and opens in a tube beneath the strut (pterygoid plate) between the pterygoid canal and the foramen ovale (fig. 8A); it sometimes lacks the dorsal wall and appears absent. There are no sphenopalatine vacuities. Auditory bullae not inflated, nearly flat in lateral profile, and markedly smaller than those of most other echimyines (figs. 7D, 8B); meatus rim pointed outward. Postglenoid foramen slitlike and enclosed below by a crest of petrosal. The ridges and grooves posterior to the end of the incisive foramina terminate anterior to the premolars, with shallow or absent extensions onto the palate between the toothrows (fig. 8A, B). Viewed dorsally, the interparietal is an evenwidth, narrow strip bowed smoothly posteriorly at the parietal suture (fig. 9D). Coronoid process of the mandible slender and recurved, above a shallow sigmoid notch (fig. 7D). Masseteric crest strongly developed and curved upward anteriorly for about half the width of the mandible; spine of the condyloid ridge on the interior of the mandible does not extend much more than halfway up the condyloid process, except on a specimen from southern Peru. Maxillary toothrows short, 7.3–10.75 mm (table 6); incisors bone white; roots of the upper incisors lie outside the maxillary base of the zygoma. Lower premolar quadralophodont, comprised of a curved anteroloph and a straight metolophid that join with wear into a D-shape around a central fossettid, which joins with wear by a central mure to the lingually opening V-shaped posterior lophid pair (fig. 11B). Hypoflexids on m1 to m3 fall nearly opposite the middle of the mesolophids, such that the teeth viewed from the lingual side are nearly symmetrical, rounded M-shapes. Upper lingual and lower labial reentrant folds (hypoflexi) short and wide, about 1/4 of the width of tooth, when worn they form subcircular fossettes; labial reentrant folds are long, and of equal length (meso- and metaflexi), such that when worn to a mure in M1 and M2, the mure is on the far lingual edge of the tooth (fig. 11B). Mandibular cheekteeth all with curved posterior borders. M3 trilophodont in all specimens examined, with the third loph much reduced (fig. 11C). Worn lower molars nearly circular, somewhat resembling those of Isothrix (fig. 11C). COMPARISONS WITH OTHER GENERA: Leiuromys is most easily confused externally with the distantly related taxa Mesomys species and Phyllomys pattoni. Species of Mesomys all have conspicuously hairy tails with brown, outward-curling hairs and a slight-to-prominent pencillike tapering at tip (Patton et al., 2000). In Mesomys the buff or whitish ventral pelage field is sharply demarcated from the spiny dorsal pelage. The molar occlusal pattern has flexi/flexids compressed into narrow parallel lines that do not open on sides of the teeth. Phyllomys pattoni have rusty-tipped spines and rusty sides, and an occlusal pattern of separate laminae (Emmons et al., 2002). Pattonomys species are larger, with grizzled gray heads, sides, and feet, white lower cheeks and throat, a fuzzy tail, and a distinct molar occlusal pattern, including straight posterior edges of the lower cheekteeth (fig. 12E–G). The spines of Pattonomys species taper more sharply at the tips (fig. 4B). Makalata species have tails slightly but distinctly hairy, with banded tail hairs, and short, rectangular tail scales in even rings (fig. 5D); a uniformly ochraceous- or reddish-agouti pelage of dusky and yellowish; multibanded, spinous aristiform hairs with short, pale, hairlike tips that are not as wide or sharp as in Leiuromys (fig. 4C); rostra that are reddish anteriorly to above and behind the eyes (fig. 13C), and rusty pelage around the tail base. Phyllomys species and Echimys chrysurus have one inguinal and three lateral pairs of mammae, whereas the other genera have two lateral pairs, and if present, inguinal mammae seem obsolete (unused in parous females). All other Echimyini usually have a quadralophodont M3, but P. semivillosus can lack the fourth loph. In Makalata the lower premolars are pentalophodont (illustrated in Emmons, 2005). The lower premolar hypoconid is a rounded point in genera of the Echimys clade, while it is flat in Pattonomys and Toromys; in the latter often slightly indented (Iack-Ximenez et al., 2005: fig. 11)., Published as part of Emmons, Louise H. & Fabre, Pierre-henri, 2018, A Review of the Pattonomys / Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus, pp. 1-52 in American Museum Novitates 2018 (3894) on pages 13-24, DOI: 10.1206/3894.1, http://zenodo.org/record/5369712, {"references":["Thomas, O. 1921. New Sigmodon, Oryzomys, and Echimys from Ecuador. Annals and Magazine of Natural History (9) 7: 448 - 450.","Smithe, F. B. 1975. Naturalist's color guide. New York: American Museum of Natural History.","Patton, J. L., M. N. F. da Silva, and J. R. Malcolm. 2000. Mammals of the Rio Jurua and the evolutionary and ecological diversification of Amazonia. Bulletin of the American Museum of Natural History 244: 1 - 306.","Emmons, L. H., Y. L. R. Leite, D. Kock, and L. P. Costa. 2002. A review of the named forms of Phyllomys (Rodentia: Echimyidae) with the description of a new species from coastal Brazil. American Museum Novitates 3380: 1 - 40.","Emmons, L. H. 2005. A revision of the arboreal Echimyidae (Rodentia: Echimyidae, Echimyinae); with descriptions of two new genera. In E. A. Lacey, and P. Myers (editors), Mammalian diversification: from chromosomes to phylogeography: 247 - 309. Berkeley: University of California Press."]}

Emmons, Louise H., Fabre, Pierre-henri (2018): A Review of the Pattonomys/Toromys Clade (Rodentia: Echimyidae), with Descriptions of a New Toromys Species and a New Genus. American Museum Novitates 2018 (3894): 1-52, DOI: 10.1206/3894.1, URL: http://www.bioone.org/doi/10.1206/3894.1

Melomys burtoni from Halmahera Referred specimens Ten specimens of Melomys burtoni from Halmahera were caught near the Gemaf and Sagea villages on June 2010 and September 2013 (MZB 33533, 33537, 33539, 33540, 33225, 36359, 36360, 36361, 36362, 36363), and eight specimens were collected on the South (MZB 33538) and North Ake Jira (Sake River; MZB 33223, 33224, 33226, 33534, 33536, 33541, 33552) in January 2010, 19 km NW of Gemaf and 26 km NW of Sagea at the GPS coordinates 00°37′23.8″N, 127°55′12.6″E (South Ake Jira) and 00°38′49.9″N, 127°56′01.2″E (North Ake Jira); Ake Sake (Sake River), 5 km NW of the village of Gemaf and 12 km NW of the village of Sagea, Central Halmahera, Halmahera Island, Indonesia, at an altitude of 22 m, at the GPS coordinates 00°29′14.6″N, 127°59′15.7″E (see Figure 1). Their current altitudinal range is 0 from 250 m. Seventeen specimens are prepared as study skins with accompanying skulls. Distribution Currently, Melomys burtoni from Halmahera is known only from the south-central region of the North Moluccan island of Halmahera (Figure 1). The species has been collected in lowland forest and forest edge habitats, at elevations between 0 and 250 m. We suspect that it may occur more broadly in other regions in the tropical lowland evergreen rain forest (Whitmore et al. 1984) in Halmahera, and on neighbouring islands such as Morotai and perhaps Bacan, but little is yet known of its biology and distribution. Previous efforts to survey the rodent fauna of Halmahera and adjacent islands (specimens deposited at AM, AMNH, MZB, USNM), which have documented two native rodent species (Rattus morotaiensis and Halmaheramys bokimekot) and several non-native rodents (Rattus exulans, R. tanezumi / R. rattus, and Mus), have not yet documented Melomys in other areas of Halmahera or on Morotai or Bacan. However, rodent collecting efforts in the North Moluccas have been very limited to date. Sampling was done at higher elevation in Halmahera by us (900– 1000 m, cf. boki mekot locality Fabre et al. 2013) and also other mammalogists (Flannery 1995a,b), but M. burtoni were never collected above 250 m. aMZB 33223-26; 33533-34; 33536-41; 33552. bBMNH .3.4.10.1; AU 24391-92. cAMNH 151204-215; 151217; 151223-227. dAMNH 105754-760; 105762-771; 105866; 105964-965; 105967-969. eAMNH 154116; 154120; 154123; 154125-126; 154128-129; 154131; 154133; 154135; 154136-138; 154140-141; 154144; 154149-153. aMZB 35961; AU 31923. bBMNH .20.7.26.26-27. cBMNH .20.7.26.21-25; BMNH.20.7.26.22. dBMNH .20.7.26.19-20. eBMNH .21.2.9.2. fBMNH .21.2.9.3. Measurements See Tables 4–6. Diagnosis Melomys burtoni from Halmahera can be distinguished from congeners by the following external characters: (1) relatively small body size, e.g. as compared to M. aerosus and M. fulgens of Seram (Helgen 2003) (Tate 1951, Menzies aMZB 33224-26; 33533-34; 33536-41; 33552. 1996, see also Figures 6 and 7; Tables 5 and 6); (2) head-body length is nearly equal to the tail length (103 % of the body length, but see also Figures 6 and 7) compared to the long-tailed M. obiensis (127 % of the body length); (3) dorsal fur dark brown or red-brown with dark gray bases; (4) ventral fur pure white along mid-ventral region of underparts and grayish on the sides; (5) dorsal surfaces of the forefeet and hindfeet pale brown with scattered rufous hairs; (6) tail usually dark above and below (some specimens have a paler underside), with three hairs per tail scale; (7) brown ears relatively long and large. Craniodental features diagnostic in combination include (1) a long rostrum, topped by broad nasals and slightly convex at the premaxillary tips (Figures 5, 8 and 9); (2) a narrow interorbital region with concave borders (Figure 8); (3) large palatine foramina (Figure 10), (4) long incisive foramina that extend posteriorly to the start of the molar rows (Figures 4, 5 and 10), (5) relatively longer incisive foramina and shorter tympanic bullae compared with all other species sampled here (Figures 4, 5 and 10). Following Menzies (1996), Melomys burtoni from Halmahera belongs within the “ cervinipes division” and more specifically is a member of the M. burtoni species group that includes M. burtoni of Australia, M. lutillus and M. frigicola of New Guinea, and four insular taxa distributed both west and east of New Guinea (M. bannisteri, M. cooperae, M. howi, M. paveli). Six species (M. aerosus, M. caurinus, M. fulgens, M. obiensis, M. fraterculus, M. talaudium) from the Moluccan region should not be included in this group due to lack of data as well as their relative morphological divergence. Description Melomys burtoni from Halmahera (Figures 6, 7, and 11) is relatively small in body size (head-body length 110–141 mm) with short, dense, and soft pelage. Dorsal fur ranges from dark-brown to red-brown; the hairs are bicolored, uniformly gray to blackish at the bases (the proximal 1/4 of the length) and brightly rufous distally. Dorsal guard hairs are soft, short, and pliable, project up to 7 mm beyond the overfur, and are glistening black for most of their length with some reddish tips. The ventral coat is pure white the length of the midventral region and grayish towards the flanks. We detect major chromatic differences between the pelage of an old adult (n = 1), and other adults (n = 15). The old individual MZB 53533 has an overall darker coat and large body size, but is molecularly identical to all other M. burtoni specimens from Halmahera. The studied juvenile (n = 1) display a finer and duller colour tone and a more greyish fur compared to the adults. The snout of Melomys burtoni from Halmahera is reddish-brown to yellowish above, and white on the sides and underside. The rhinarium and lips are unpigmented. The eyes are small (diameter about 5 mm). Vibrissae are dark gray. Mystacial and superciliary vibrissae are moderately long, extending slightly beyond the ears. The pinnae are relatively small and brown, with a sparse covering of reddish brown hairs on both the inner and outer surfaces. The tail is nearly equal in length to the combined head and body length (only exceeding HBL in the specimen MZB 33533) (Tables 4 and 6). The tail is dark brown to blackish on the entire surface, except for the proximal portion (10–25 mm), which is unpigmented in some specimens (MZB 33533, 33552). Tail scales are smooth and less raised compared to those in species of the M. rufescens Division (Menzies 1996), and moderately large (14–15 rows/cm). There are three hairs per tail scale (each 1–2 mm long), which is characteristic of the M. cervinipes Division, as described by Menzies (1996). Tail hairs are dark brown. Dorsal surfaces of the forefeet and hindfeet are pale brown or unpigmented (Figures 6) and sparsely covered with short reddish or brownish hairs on top of the metapodials and laterally on the digits. At the base of each claw, long white hairs form ungual tufts. The pollex is very short and bears a flat nail. The other front digits are longer and broader with curved, sharp, white claws. Each palmar surface is hairless, unpigmented, and displays three interdigital pads and two large metacarpal pads. The hindfeet of Melomys burtoni from Halmahera are shorter than those of M. obiensis and those of M. frigicola (Tables 4 and 5). Digits are elongated and broad, and each bears a pale claw that is short, sharp, and recurved. The hallux extends past the middle part of the medial digits. The 5th digit is slightly shorter than the medial digits and has a large associated interdigital pad. The plantar surface is naked and unpigmented, with four large interdigital pads, an elongate thenar pad, and a very small hypothenar pad. As in all Melomys species, it has only two pairs of inguinal teats (0 + 2 = 4). The skull of Melomys burtoni from Halmahera (Figure 11) is smaller than most Moluccan species of Melomys and similar to M. burtoni and allies in most dimensions (Figures 4, 8–10). In dorsal view, the rostrum is narrow and short (Figure 11). The interorbital region is moderately narrow and defined by concave borders. The postorbital region is elongate and the inflated braincase has a rounded outline. The temporal ridges that extend from the supraorbital region to the dorsolateral surfaces of the braincase are slightly developed. Robust zygomatic arches bow out from rostrum and braincase. In lateral view (Figures 9 and 11), the dorsal profile of the skull is broadly convex from the slightly depressed nasal tips to the top of the frontals, straight over the frontals and parietals to the apex of the domed braincase, and sloping abruptly downward over the length of the interparietal. Nasal bones are short with sloping sides that give an angular shape to the tip of the rostrum. The posterior portion of the nasals are flat with nearly no concavity at the junction with the proximal end of the frontal. The zygomatic plate is narrower and the zygomatic notch shallower than in other Moluccan Melomys. The dorsal root and anterior portion of the zygomatic arch are slightly broadened. The squamosal root of the arch is positioned at the medial portion of the braincase. A subsquamosal fenestra is not present. Each auditory bulla is small (Figures 4, 5, 10, and 11) and separated from the squamosal by a small postglenoid vacuity and from the alisphenoid by a narrow middle lacerate foramen. The lambdoid crest is poorly developed. The ventral view of the skull (Figures 10 and 11) shows long incisive foramina (Figures 4 and 5) that project posteriorly beyond the anterior margins of the first molars. The palate is narrow (BPPM1 and BPPM3) and short (LBP). Each palatine foramen is large, positioned between the anterior and posterior roots of the M2, and opens anteriorly into a well-defined palatal sulcus that runs forward to the posterolateral margin of the incisive foramina. The mesopterygoid fossa is moderately broad. The lateral pterygoid fossae are shallow and contain large sphenopterygoid vacuities. The skull exhibits bony landmarks consistent with the primitive pattern of cephalic arterial circulation found in most murine rodents (Musser and Heaney 1992), as in other Melomys (Menzies 1996). The upper and lower molars of Melomys burtoni from Halmahera are brachydont and broader compared to M. lutillus from Papua and M. burtoni from Australia (Figures 12 and 13; Tables 6 and 7). Each molar overlaps with its neighbour. The first upper and lower molars form nearly half of the entire toothrow, the second about onethird of the row, with the last molar much smaller than the others (Figures 12 and 13). The transverse laminae of the upper molars (rows 1 and 2 of M1 and row 2 of M2) display a posteriorly oriented, angular U-shape with a typically acuspidate chevron pattern (characteristic of Melomys, Paramelomys and Uromys). The first row is formed by a medium-sized cusp t1 slightly separated from a wide central cusp t2, which in turn is broadly fused with a smaller cusp t3. The second row is composed of a large backward-directed lingual cusp t4 that is attached to a wide and narrow central cusp t5 and a large labial cusp t6, which is smaller than t4. The third row is formed by a large, angular cusp t8 joined to a cusp t9 that projects straight to the labial side. The second molar consists of two laminae with a medium-sized cusp t1. The first lower molar has an oblong outline and three rows of cusps (Figure 13). The anterior 2/3 of the molar is formed by small anterolingual and anterolabial cusps that are slightly separated in unworn teeth from the second cusp row (metaconid, protoconid). The metaconid and protoconid are large, with a triangular shape, and fused in specimens displaying worn teeth (e.g. MZB 33533; Figure 13). A large entoconid and hypoconid comprise the third cusp row, behind which is a broad but anteroposteriorly narrow posterior cingulum. The subsquare second lower molar has two cusp rows and a posterior cingulum. The pattern and shape of the lamina are similar in cusp shape and pattern to first lower molar. There is a small anterolabial cusplet on the anterolabial margin of the second lower molar close to the protoconid. The small third lower molar displays a rounded shape and a simple occlusal surface with two laminar cusp rows and a posterior cingulum. The molars of Melomys burtoni from Halmahera have multiple roots, which were identified by a µCT scanner. Below each first upper molar, we found one large anterior root, two smaller lingual anchors, one labial root and one large posterior root. Four roots anchor each second molar, there are two anterior roots and two posterior anchor. Three roots anchor each third molar, there are two anterior roots and one posterior anchor. One anterior and posterior roots coupled with smaller labial and lingual anchors attach to each first lower molar. The second lower molars are anchored by two anterior and two posterior roots. The third lower molars are anchored by two anterior and one posterior roots. The alveoli patterns of upper and lower molar roots seen in Melomys burtoni from Halmahera are similar to the patterns found for M. lutillus from Papua and M. burtoni from Australia. Each dentary of Melomys burtoni from Halmahera is moderately long and stocky (Figure 11), with a short but stout coronoid process, robust condylar process and strong angular process. There is a concave curvature of the posterior margin of the dentary. The capsular process forms a prominent ridge parallel to the coronoid and anterior part of the condylar process. A prominent masseteric crest extends between the ventral margin and the labial part of the dentary just before the mental foramen. The mandibular foramen is located above a narrow ridge that runs caudally from the molar platform to the base of the condylar process., Published as part of Fabre, Pierre-Henri, Fitriana, Yuli S., Semiadi, Gono, Pagès, Marie, Aplin, Ken, Supriatna, Nanang & Helgen, Kristofer M., 2018, New record of Melomys burtoni (Mammalia, Rodentia, Murinae) from Halmahera (North Moluccas, Indonesia): a review of Moluccan Melomys, pp. 218-247 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 82 (3) on pages 228-237, DOI: 10.1515/mammalia-2016-0137, http://zenodo.org/record/7837666, {"references":["Whitmore, T. C. 1984. Tropical rain forests of the Far East. Second edition. Oxford University Press, Oxford.","Fabre, P. - H., M. Pages, G. G. Musser, Y. S. Fitriana, J. Fjeldsa, A. Jennings, K. A. JOnsson, J. Kennedy, J. Michaux, G. Semiadi, N. Supriatna and K. M. Helgen. 2013. A new genus of rodent from Wallacea (Rodentia: Muridae: Murinae: Rattini), and its implication for biogeography and Indo-Pacific Rattini systematics. Zool. J. Linnean Soc. 169: 408 - 447.","Flannery, T. F. 1995 a. Mammals of the south-west Pacific and Moluccan Islands. Reed Books, Chatswood, New South Wales, Australia.","Helgen, K. M. 2003. A review of the rodent fauna of Seram, Moluccas, with the description of a new subspecies of mosaic-tailed rat, MelomyS rUfeScenS paVeli. J. Zool. (London) 261: 165 - 172.","Tate, G. H. H. 1951. Results of the Archbold Expeditions. No. 65. The rodents of Australia and New Guinea. Bull. Am. Mus. Nat. Hist. 97: 183 - 430.","Menzies, J. I. 1996. A systematic revision of MelomyS (Rodentia: Muridae) of New Guinea. Aust. J. Zool. 44: 367 - 426.","Musser, G. G. and L. R. Heaney. 1992. Philippine rodents: definitions of TarSomyS and LimnomyS plus a preliminary assessment of phylogenetic patterns among native Philippine murines (Murinae, Muridae). Bull. Am. Mus. Nat. Hist. 211: 1 - 138."]}

Melomys Thomas 1922a Type species Melomys rufescens (Alston 1877), by original designation. Diagnosis p = p-value. (cp Melomys is a murid genus classified in the subfamily Murinae (sensu Musser and Carleton 2005), within the tribe Hydromyini (Aplin and Helgen 2010), in the “ Uromys Legend similar to that of Table 1. The symbol “:” indicates tested interactions between explanatory variables. (ap < 0.05, bp Paramelomys (cf. Menzies 1996), (6) narrow and deep pterygoid fossae, (7) lateral extension of the parietal bone, (8) a wide alisphenoid strut that is fused to the alisphenoid canal and foramen ovale, (9) very well-developed lambdoid crests which curl forward over the squamoso-mastoid foramen, (10) a carotid arterial circulation that is primitive for murine rodents, as opposed to the derived pattern seen in Mammelomys (11) a large M 3 compared to Paramelomys, with an antero-internal cusp always present, and (12) only two pairs of inguinal teats (mammary formula 0 + 2 = 4). The Melomys population from Halmahera is characterized by small to medium body size in the genus (Weight range = 110–141 g), and has all of the diagnostic characters listed above. Content and distribution. The genus Melomys contains 23 species (Musser and Carleton 2005). It is distributed in eastern and northern Australia, throughout New Guinea, and in adjacent archipelagos, from the Moluccas in the west to the Solomon Islands in the east (Helgen 2003, Breed and Aplin 2008)., Published as part of Fabre, Pierre-Henri, Fitriana, Yuli S., Semiadi, Gono, Pagès, Marie, Aplin, Ken, Supriatna, Nanang & Helgen, Kristofer M., 2018, New record of Melomys burtoni (Mammalia, Rodentia, Murinae) from Halmahera (North Moluccas, Indonesia): a review of Moluccan Melomys, pp. 218-247 in Mammalia (Warsaw, Poland) (Warsaw, Poland) 82 (3) on pages 227-228, DOI: 10.1515/mammalia-2016-0137, http://zenodo.org/record/7837666, {"references":["Thomas, O. 1922 a. A subdivision of the genus UromyS. Ann. Mag. Nat. Hist. (series 9) 9: 260 - 261.","Alston, E. R. 1877. On the rodents and marsupials collected by the Rev. G. Brown in Duke of York Island, New Britain and New Ireland. Proceedings of the Zoological Society of London 1877: 123 - 127.","Musser, G. G. and M. D. Carleton. 2005. Superfamily Muroidea.","Aplin, K. P., Helgen, K. M. 2010. Quaternary murid rodents of Timor. Part I: New material of CoryphomyS bUehleri Schaub, 1937, and description of a second species of the genus. Bull. Am. Mus. Nat. Hist. 341: 1 - 80.","Menzies, J. I. 1996. A systematic revision of MelomyS (Rodentia: Muridae) of New Guinea. Aust. J. Zool. 44: 367 - 426.","Helgen, K. M. 2003. A review of the rodent fauna of Seram, Moluccas, with the description of a new subspecies of mosaic-tailed rat, MelomyS rUfeScenS paVeli. J. Zool. (London) 261: 165 - 172.","Breed, W. G. and K. P. Aplin. 2008. The ' UromyS Group': Papuan"]}

Tyto almae, sp. nov. Seram Masked Owl Holotype. Museum Zoologicum Bogoriense (MZB), Cibinong, Indonesia, study skin voucher: MZB 33.231, female (ovary 17 × 5 mm), Mount Binaiya, at Waensela above Kanikeh Village, Manusela National Park, Seram, Maluku Province, Indonesia, elevation 1,350 m, 3 °08.668’S, 129 ° 28.434 ’E, mist-netted on 10 February 2012; prepared by Knud Andreas Jønsson, field catalogue number KAJ 1 - 10.02.12. Measurements of holotype. Wing length (chord): 252 mm; tarsus: 63.6 mm; tail: 116 mm; bill from base of skull: 37.4 mm; bill height at base: 16.1 mm; bill width: 13.7 mm; weight: 540 g. Description of holotype. Facial area light pinkish cinnamon, more dusky in the eye-pit, with short feathers that do not cover the auricular troughs (probably as a consequence of heavy moult), with a ruff of orange tawny feathers with dark brown tips bordering the facial area outside the auricular troughs. Crown, nape, mantle and wing coverts ochraceous orange, each feather with a short white stripe embedded within dusky-mottled feather-tips; lower back to upper tail-coverts with more spindle-shaped and pale buffy spots within the dark feather-tips. Greater wing-coverts and remiges ochraceous orange with well-spaced fuscous bars and only a few darker mottles between bars; upper side of rectrices similar, ochraceous tawny with five exposed fuscous bars, each 5–8 mm wide, and with a few dusky mottles or stipples within the broader ochraceous zones. The entire extent of the underparts, including wing linings, yellow-ocher with whitish basal parts of the feathers, and small, roundish subterminal spots on most feathers of the breast and belly; tarsi with yellow-ocher feathers all the way to the base of the toes. Irides dark brown, bill pale horn, feet pinkish drab with pale grey talons. Diagnosis. Head large and round with a heart-shaped facial area, tail short, and legs rather long as is typical of masked-owls of the genus Tyto (Bruce 1999). The general colouration, size and geographical location initially led us to believe that it belonged within Tyto sororcula. Generally similar to both forms of Tyto sororcula, but differs by having yellow ochre instead of white or whitish underparts. Also shows a unique pattern on crown and upper back, with a short white or buff shaft stripe within the dusky terminal part of each feather, instead of two roundish, or bar- or heart-shaped white spots per feather, as in most Tyto species. Additionally, feathers of the mid- and lower back have only a limited amount of light (buffy) mottles within the dusky terminal parts, unlike in T. s. cayelii and, notably, T. s. sororcula. In the latter subspecies, distal parts of the dorsal feathers are densely mottled with dusky and white, and there is also a broad mottled zone inside each of the dark bars, giving a rather ‘peppered’ greyish appearance to much of the upperparts. The tail of T. almae is extensively golden brown with broad dark bars, showing no pale mottling, and only a trace of dark mottling between the bars. This is unlike the more extensive dark mottling found in both forms of T. sororcula, which additionally shows pale mottling. Thus, with more extensively golden brown and distinctly barred wings and tail, T. almae is phenotypically quite distinct from the two currently recognized forms of Tyto sororcula. Furthermore, T. almae has longer and completely feathered tarsi, similar to T. s. cayelii, whereas the lower tarsi of T. s. sororcula have short bristle-like feathers only. Etymology. The specific epithet honours Alma Jønsson, daughter of the senior author, acknowledging that she had to be without her father while he was out exploring. Conservation. Both known records of Tyto owls from Seram are from within the protected Manusela National Park. Manusela National Park covers about 10 % of Seram (17,100 km 2). This is one of the only areas in Seram that is visited with any regularity by ornithologists (Bowler & Taylor 1989, 1993; Marsden et al. 1997; Rheindt & Hutchinson 2007 a,b) and other individuals who would recognize the scientific significance of a Tyto owl on Seram. The natural vegetation of Seram is tropical lowland evergreen and semi-evergreen rain forest, with tropical montane rain forest above ~ 800 m. Seram is still well-forested, although the low-lying areas where the human population is concentrated along the coast and in the west have been cleared (BirdLife International 2012). It is very problematic to assess the threat status of this species as so little is known about its distribution, status and habitat requirements. The little information available suggests that T. almae is not threatened at present, but lives at rather low population densities within montane and perhaps lowland rain forest on Seram. Timber extraction and bird trapping for the wild bird trade are known threats to other forest bird species of Seram (Marsden 1998; Chan et al. 2004; BirdLife International 2012). The most likely threat to Tyto almae is disappearance of its forest habitat due to anthropogenic activities. The existence of Manusela National Park may be of high importance for the longterm conservation of this species.

Tyto almae, sp. nov. Seram Masked Owl Holotype. Museum Zoologicum Bogoriense (MZB), Cibinong, Indonesia, study skin voucher: MZB 33.231, female (ovary 17 �� 5 mm), Mount Binaiya, at Waensela above Kanikeh Village, Manusela National Park, Seram, Maluku Province, Indonesia, elevation 1,350 m, 3 ��08.668���S, 129 �� 28.434 ���E, mist-netted on 10 February 2012; prepared by Knud Andreas J��nsson, field catalogue number KAJ 1 - 10.02.12. Measurements of holotype. Wing length (chord): 252 mm; tarsus: 63.6 mm; tail: 116 mm; bill from base of skull: 37.4 mm; bill height at base: 16.1 mm; bill width: 13.7 mm; weight: 540 g. Description of holotype. Facial area light pinkish cinnamon, more dusky in the eye-pit, with short feathers that do not cover the auricular troughs (probably as a consequence of heavy moult), with a ruff of orange tawny feathers with dark brown tips bordering the facial area outside the auricular troughs. Crown, nape, mantle and wing coverts ochraceous orange, each feather with a short white stripe embedded within dusky-mottled feather-tips; lower back to upper tail-coverts with more spindle-shaped and pale buffy spots within the dark feather-tips. Greater wing-coverts and remiges ochraceous orange with well-spaced fuscous bars and only a few darker mottles between bars; upper side of rectrices similar, ochraceous tawny with five exposed fuscous bars, each 5���8 mm wide, and with a few dusky mottles or stipples within the broader ochraceous zones. The entire extent of the underparts, including wing linings, yellow-ocher with whitish basal parts of the feathers, and small, roundish subterminal spots on most feathers of the breast and belly; tarsi with yellow-ocher feathers all the way to the base of the toes. Irides dark brown, bill pale horn, feet pinkish drab with pale grey talons. Diagnosis. Head large and round with a heart-shaped facial area, tail short, and legs rather long as is typical of masked-owls of the genus Tyto (Bruce 1999). The general colouration, size and geographical location initially led us to believe that it belonged within Tyto sororcula. Generally similar to both forms of Tyto sororcula, but differs by having yellow ochre instead of white or whitish underparts. Also shows a unique pattern on crown and upper back, with a short white or buff shaft stripe within the dusky terminal part of each feather, instead of two roundish, or bar- or heart-shaped white spots per feather, as in most Tyto species. Additionally, feathers of the mid- and lower back have only a limited amount of light (buffy) mottles within the dusky terminal parts, unlike in T. s. cayelii and, notably, T. s. sororcula. In the latter subspecies, distal parts of the dorsal feathers are densely mottled with dusky and white, and there is also a broad mottled zone inside each of the dark bars, giving a rather ���peppered��� greyish appearance to much of the upperparts. The tail of T. almae is extensively golden brown with broad dark bars, showing no pale mottling, and only a trace of dark mottling between the bars. This is unlike the more extensive dark mottling found in both forms of T. sororcula, which additionally shows pale mottling. Thus, with more extensively golden brown and distinctly barred wings and tail, T. almae is phenotypically quite distinct from the two currently recognized forms of Tyto sororcula. Furthermore, T. almae has longer and completely feathered tarsi, similar to T. s. cayelii, whereas the lower tarsi of T. s. sororcula have short bristle-like feathers only. Etymology. The specific epithet honours Alma J��nsson, daughter of the senior author, acknowledging that she had to be without her father while he was out exploring. Conservation. Both known records of Tyto owls from Seram are from within the protected Manusela National Park. Manusela National Park covers about 10 % of Seram (17,100 km 2). This is one of the only areas in Seram that is visited with any regularity by ornithologists (Bowler & Taylor 1989, 1993; Marsden et al. 1997; Rheindt & Hutchinson 2007 a,b) and other individuals who would recognize the scientific significance of a Tyto owl on Seram. The natural vegetation of Seram is tropical lowland evergreen and semi-evergreen rain forest, with tropical montane rain forest above ~ 800 m. Seram is still well-forested, although the low-lying areas where the human population is concentrated along the coast and in the west have been cleared (BirdLife International 2012). It is very problematic to assess the threat status of this species as so little is known about its distribution, status and habitat requirements. The little information available suggests that T. almae is not threatened at present, but lives at rather low population densities within montane and perhaps lowland rain forest on Seram. Timber extraction and bird trapping for the wild bird trade are known threats to other forest bird species of Seram (Marsden 1998; Chan et al. 2004; BirdLife International 2012). The most likely threat to Tyto almae is disappearance of its forest habitat due to anthropogenic activities. The existence of Manusela National Park may be of high importance for the longterm conservation of this species., Published as part of J��nsson, Knud Andreas, Poulsen, Michael K��ie, Haryoko, Tri, Reeve, Andrew Hart & Fabre, Pierre-Henri, 2013, A new species of masked-owl (Aves: Strigiformes: Tytonidae) from Seram, Indonesia, pp. 51-61 in Zootaxa 3635 (1) on pages 55-57, DOI: 10.11646/zootaxa.3635.1.5, http://zenodo.org/record/283524

Jønsson, Knud Andreas, Poulsen, Michael Køie, Haryoko, Tri, Reeve, Andrew Hart, Fabre, Pierre-Henri (2013): A new species of masked-owl (Aves: Strigiformes: Tytonidae) from Seram, Indonesia. Zootaxa 3635 (1): 51-61, DOI: http://dx.doi.org/10.11646/zootaxa.3635.1.5

International audience; We describe Halmaheramys bokimekot Fabre, Pages, Musser, Fitriana, Semiadi & Helgengen.etsp.nov., a new genus and species of murine rodent from the North Moluccas, and study its phylogenetic placement using both molecular and morphological data. We generated a densely sampled mitochondrial and nuclear DNA data set that included most genera of Indo-Pacific Murinae, and used probabilistic methodologies to infer their phylogenetic relationships. To reconstruct their biogeographical history, we first dated the topology and then used a Lagrange analysis to infer ancestral geographic areas. Finally, we combined the ancestral area reconstructions with temporal information to compare patterns of murine colonization among Indo-Pacific archipelagos. We provide a new and comprehensive molecular phylogenetic reconstruction for Indo-Pacific Murinae, with a focus on the Rattus division. Using previous results and those presented in this study, we define a new Indo-Pacific group within the Rattus division, composed of Bullimus, Bunomys, Paruromys, Halmaheramys, Sundamys, and Taeromys. Our phylogenetic reconstructions revealed a relatively recent diversification from the Middle Miocene to Plio-Pleistocene associated with several major dispersal events. We identified two independent Indo-Pacific dispersal events from both western and eastern Indo-Pacific archipelagos to the isolated island of Halmahera, which led to the speciations of H.bokimekotgen.etsp.nov. and Rattus morotaiensis Kellogg, 1945. We propose that a Middle Miocene collision between the Halmahera and Sangihe arcs may have been responsible for the arrival of the ancestor of Halmaheramys to eastern Wallacea. Halmaheramys bokimekotgen. et sp.nov. is described in detail, and its systematics and biogeography are documented and illustrated.(c) 2013 The Linnean Society of London