Subgenus TRINOMYS Thomas

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Genotype.Echimys albispinus Is. Geoffroy Saint-Hilaire, 1838; by original designation.

Trinomys Thomas, July 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):140 (orig. descr.); Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):401; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:115.

Fig. 99. Fig. 99.Map showing the geographic ranges of the subspecies of three species of the subgenus Trinomys.

General characters.—Pelage of upper parts with lanceolate and, sometimes, clavate aristiforms extending over most of rump and onto thighs; tail 86 to 103 per cent of length of head and body; tail sometimes white-tipped and sometimes penicillate; skull small, with ridges moderately developed; supraorbital ridges involving no part of parietals; infraorbital foramen with no separate groove for transmission of nerve; mesopterygoid fossa extending forward to level of second or first molars; incisors opisthodont, orthodont or proodont; molariform teeth, in occlusal view, with main fold large and usually reaching opposite wall; no counterfold anterior to main fold in upper molariform teeth and usually no counterfold posterior to main fold in lower molariform teeth; premolars larger than first molars, first molars larger than second molars and second molars larger than third molars; four molariform teeth of nearly equal size in some animals.

Thomas (1921:140) erected the subgenus Trinomys, including in it the species albispinus and setosus and stated that "the primary distinction between these [Trinomys and Proechimys] lies in the number of laminae present in the cheekteeth—four in Proechimys, three in Trinomys." The distinction is valueless as a subgeneric character, not only because the character is not constant in the species in the subgenus but also because there is subspecific variation in number of laminae in the cheekteeth. Proechimys albispinus, however, shares with three other species common characters, as listed above, and the name Trinomys will, therefore, apply to this group of species, since Proechimys albispinus is the genotype.

Proechimys dimidiatus(GÜnther)

Echimys dimidiatus GÜnther, 1 April 1877, Proc. Zool. Soc. London, 1876(4):747.

Proechimys dimidiatus Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20):264; Ribeiro, 1905, Arch. Mus. Nac. Rio de Janeiro, 13:187; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):141; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.

Type locality.—Unknown; probably southwestern Rio de Janeiro, Brazil (see Remarks). Type: British Museum (Nat. Hist.), no. 51.7.21.24; presented by Lord Derby.

Range.—Rio de Janeiro, from the southern limit of the state northward to and including the Distrito Federal.

General characters.—Size large; tail averaging 80 per cent of head and body; aristiforms narrow and soft (0.4 to 0.5 mm wide), imparting a non-spiny character to the pelage; general color of upper parts Ochraceous-Buff, finely lined with blackish brown, gradually becoming lighter on sides; ventral surface of body and inner sides of legs white; feet dorsally white but with a sepia-colored stripe along outer margin; tail brownish-black above and white below, but white sometimes extended to upper side in distal part; skull broad with no conspicuous ridges; jugals deep with transverse ridge usually conspicuous; postorbital process of zygoma involving only squamosal; incisive foramen short and wide posteriorly; vomerine sheath complete in 95 per cent of specimens and with maxillary part thick; posterior palatine foramina at plane of first molars or slightly anterior to them; bullae moderately developed; in juvenal specimens, each upper molariform tooth with three counterfolds, but posteriormost counterfold small; in adult specimens, posteriormost counterfold disappearing in 50 per cent of fourth premolars and first molars, in 20 per cent of second molars, and in 15 per cent of third molars; lower molariform teeth with two counterfolds in almost every juvenal specimen, this number, in adult animals, decreasing in m3 to one in 20 per cent of specimens but rarely being reduced in other teeth.

General characters.—Aristiforms soft and narrow, ranging from 15 to 19 mm in total length and 0.4 to 0.5 mm in maximum width; pelage generally non-spiny and not harsh; length of tail ranging from 20 per cent shorter than head and body to as long as, or slightly longer than, head and body; ears rather small (23 to 25 mm).

Figs. 100-103. Figs. 100-103.Proechimys dimidiatus, male, MN no. 5452, Tijuca. × 1.

Color.—General color of back and sides results from uniform mixture of black distal parts of aristiforms with Ochraceous-Buff of subapical zone of setiforms. Dorsally, from nose caudad to rump, mixture appears brownish-black, lined with Ochraceous-Buff; toward sides, amount of Ochraceous-Buff gradually increases and resultant color is much lighter brown than on back. On outer parts of arms and legs, color turns gradually to sepia toward distal parts and finally to uniform sepia on wrists and ankles, this color extending to outer dorsal parts of hands and feet; on ankles, sepia forms complete ring, as usual in the genus. Tail blackish-brown on upper parts, this stripe gradually tapering toward tip where dark brown hairs form small pencil; white of under side of tail sometimes seen also entirely around distal part, short of tip which remains dark brown. Ventral surfaces wholly white, from upper lips caudad including inner surfaces of legs.

Hairs.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip that has long, fine filament; total length 16 to 19 mm; maximum width 0.5 mm. On outer thigh whitish basally, gradually blackening toward tip; some with Ochraceous-Buff, subapical zone; total length 13 to 15 mm; maximum width 0.25 mm. Setiforms on middorsal region: Whitish on basal half, gradually blackening toward tip, but interrupted by Ochraceous-Buff, subapical zone; some with Light Ochraceous-Buff, subapical zone and short, blackish zone on tip; total length 12 to 14 mm; maximum width 0.02 mm. Setiforms on outer thighs: Whitish on basal half, then gradually becoming gray on middle part and finally Light Ochraceous-Buff on distal third, or with tip blackish and Ochraceous-Buff, subapical zone.

Skull.—Elongate and broad with no conspicuous crests; rostrum rather stout; jugals deep with transverse crest usually well-developed; zygomatic postorbital process conspicuous and formed entirely of squamosal; incisive foramen short and wide posteriorly; vomerine sheath complete in great majority of specimens, its maxillary part wide and strong; posterior palatine foramina on plane with front of M1 or slightly farther forward; bullae rather small and elongate.

Teeth.—P4 with three secondary folds in all juvenal specimens, but posteriormost fold small and disappearing in 50 per cent of adult specimens; M1 with 3 outer folds in juveniles and also disappearing in 50 per cent of adults; M2 with three outer folds in juveniles, but only 20 per cent remaining in adults; M3 with 3 outer folds in 50 per cent of juveniles, decreasing to 15 per cent in adults. Lower molariform teeth: p4 with 2 secondary folds; m1 with 2 secondary folds in 90 per cent of adults and in all juveniles; m2 with 2 secondary folds in 98 per cent of adults and in all juveniles; m3 with 2 secondary folds in 81 per cent of adults, remaining percentage with only one counterfold, and with 2 secondary folds in all juveniles.

Remarks.—Samples studied of P. dimidiatus are notably uniform throughout the geographic range of the species. The few biotypes detected seemed unworthy of subspecific rank.

In discussing the type locality of the species, Thomas (1921:141) states: "We know that its donor did obtain a number of specimens from Rio Janeiro, and the skull agrees so closely with those of two examples from Itatiaia, near to the Rio-Minas frontier, collected and presented by Prof. J. P. Hill, that I have no hesitation in referring the latter to GÜnther's species."

Specimens examined.—Total number, 211 (MN), from Brazil as follows: Rio de Janeiro; Parati, Pedra Branca (400 m.), 113; Mangaratiba, Fazenda do RubiÃo (750 m.), 3; Fazenda do Tenente (700 m.), 4; Fazenda da Lapa (450 m.), 13; TeresÓpolis, Fazenda Guinle (960 m.), 61; Nova IguassÚ, Barro Branco (20 m.), 16; Distrito Federal, Tijuca, 1.

Additional records.—Rio de Janeiro, Itatiaia (Thomas, 1921:141); Rio de Janeiro, Zona da mata, Mont-Serrat, Serra do Itatiaia (Ribeiro, 1905:187).

General characters.—Size large; tail long; aristiforms generally wide and stiff; general color on upper parts and sides a combination of blackish from tips of aristiforms with cinnamon ground color from subapical zones of setiforms; darker band on middorsal line; differentiated light-colored aristiforms conspicuous on outer sides of thighs and rump; usually rufous tint on neck and postauricular region; underparts white; tail with white tip, usually accentuated by white brush; feet white on dorsal surface; hind feet slightly darker on outer sides; skull elongate and smooth; jugals wide dorso-ventrally; incisive foramen elongate; upper molariform teeth usually with one to five counterfolds, number varying with subspecies; lower premolar always with two counterfolds and lower molars always with one or two counterfolds.

Remarks.—As a whole, the samples of the populations of the species do not afford a satisfactory record of the distribution; my concept of the group may be changed when further collections are made in localities geographically intermediate between those from which specimens now are known. If some of the forms prove to be physiologically isolated, they may deserve treatment as full species according to the conventional standards of systematic zoology. P. panema, for example, does not seem to be geographically isolated from P. gratiosus. P. denigratus, at the northernmost known occurrence of the species, actually represents a striking jump in the cline, although collections from intermediate regions may provide intermediate structural stages. Further collecting may also prove that the southern form, P. iheringi iheringi, is completely isolated from the rest of the group. However, these samples are certainly more related to each other than any one of them is to that of the other species found in the same range, namely P. dimidiatus, and all the forms in question, therefore, seem best arranged as subspecies of one full species. A clinal variation certainly exists among these forms and the most striking differences correspond to larger geographical distances.

Figs. 104-109.

Fig. 104.Proechimys iheringi iheringi, female, MN no. 6453, Ilha de SÃo SebastiÃo. × 1.

Fig. 105.Proechimys iheringi bonafidei, male, MN no. 6183, Fazenda BÕa FÉ. Type. × 1.

Fig. 106.Proechimys iheringi gratiosus, male, MN no. 4024, Floresta da Caixa Dagua. Type. × 1.

Fig. 107.Proechimys iheringi panema, female, MN no. 8288, Campinho. Type. × 1.

Fig. 108.Proechimys iheringi denigratus, male, MN no. 8500, Mata do RibeirÃo da Fortuna. Type. × 1.

Fig. 109.Proechimys iheringi paratus, female, MN no. 4012, Floresta da Capela de SÃo Braz. Type. × 1.

Figs. 110-115.

Fig. 110.Proechimys iheringi iheringi, female, MN no. 6453, Ilha de SÃo SebastiÃo. × 1.

Fig. 111.Proechimys iheringi bonafidei, male, MN no. 6183, Fazenda BÕa FÉ. Type. × 1.

Fig. 112.Proechimys iheringi gratiosus, male, MN no. 4024, Floresta da Caixa Dagua. Type. × 1.

Fig. 113.Proechimys iheringi panema, female, MN no. 8288, Campinho. Type. × 1.

Fig. 114.Proechimys iheringi denigratus, male, MN no. 8500, Mata do RibeirÃo da Fortuna. Type. × 1.

Fig. 115.Proechimys iheringi paratus, female, MN no. 4012, Floresta da Capela de SÃo Braz. Type. × 1.

Figs. 116-127.

Figs. 116, 117.Proechimys iheringi iheringi, female, MN no. 6453, Ilha de SÃo SebastiÃo. × 1.

Figs. 118, 119.Proechimys iheringi bonafidei, male, MN no. 6183, Fazenda BÕa FÉ. Type. × 1.

Figs. 120, 121.Proechimys iheringi gratiosus, male, MN no. 4024, Floresta da Caixa Dagua. Type. × 1.

Figs. 122, 123.Proechimys iheringi panema, female, MN no. 8288, Campinho. Type. × 1.

Figs. 124, 125.Proechimys iheringi denigratus, male, MN no. 8500, Mata do RibeirÃo da Fortuna. Type. × 1.

Figs. 126, 127.Proechimys iheringi paratus, female, MN no. 4012, Floresta da Capela de SÃo Braz. Type. × 1.

Proechimys iheringi iheringiThomas

Proechimys iheringi Thomas, August, 1911, Ann. Mag. Nat. Hist., 8 (ser. 8):252 (orig. descr.); Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):141; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.

Type locality.—Island of SÃo SebastiÃo (off SÃo Paulo), Formosa, SÃo Paulo, Brazil. Type: British Museum (Nat. Hist.), no. 2.8.25.5, adult male, presented by the SÃo Paulo Museum.

Range.—Littoral and islands of SÃo Paulo and Rio de Janeiro.

Diagnosis.—Aristiforms narrow; tail shorter than head and body; setiforms Cinnamon-Buff; incisive foramen short; vomerine sheath complete; upper molariform teeth with two or three counterfolds; lower molariform teeth with two counterfolds, rarely one in m3.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length 18 to 23 mm; maximum width, 0.6 mm. Aristiforms on outer thighs: Gray basally, blackening distally toward tip; some differentiated with Cinnamon-Buff tip. Setiforms on middorsal region: Gray basally, gradually blackening toward tip but interrupted by a Cinnamon-Buff, subapical zone 3 mm long; total length, 16 to 20 mm; maximum width, 0.06 mm. Setiforms on outer thighs: Gray basally, gradually blackening toward tip but interrupted by Cinnamon-Buff, subapical zone or with Cinnamon-Buff continuous to tip.

Skull.—Slender; bullae small and well inflated; jugal dorso-ventrally wide with transverse ridge inconspicuous; incisive foramen short, 3.5 × 2.5 mm; vomerine sheath complete; mesopterygoid fossa extending forward as far as middle parts of second molars; postorbital process of zygoma small, formed by both jugal and squamosal; posterior palatine foramina at plane of premolars; interorbital breadth narrow.

Teeth.—Upper molariform teeth with two or three counterfolds (when unworn usually three and rarely four); sometimes only one counterfold in M3 and sometimes counterfolds fused in molars. Lower molariform teeth with two counterfolds, rarely one in m3.

Comparisons.—From P. i. bonafidei and P. i. gratiosus, iheringi differs in: Incisive foramen shorter; vomerine sheath complete, instead of usually incomplete; setiforms Cinnamon-Buff, instead of Ochraceous-Buff; upper molariform teeth with two or three separate counterfolds, instead of having counterfolds fused or reduced to one or two; aristiforms narrower in iheringi than in bonafidei.

Specimens examined.—Total number, 25, from Brazil, as follows: SÃo Paulo, Formosa, Ilha de SÃo SebastiÃo, 9 (DZ 6, MN 2, MCZ 1); SÃo Paulo, Mogi das Cruzes, Alto da Serra, alt. 900 m., 2 (DZ); SÃo Paulo, Ubatuba, alt. 10 m., 4 (2 DZ, 2 MN); Rio de Janeiro, Angra dos Reis, 2 (MN); Rio de Janeiro, Angra dos Reis, Ilha Grande, 7 (5 DZ, 1 MCZ, 1 MN).

Proechimys iheringi bonafideisubspecies nova

Type locality.—Fazenda BÕa FÉ, TeresÓpolis, Rio de Janeiro, Brazil; alt. 850 meters. Type: Museu Nacional, no. 6183, adult male; collected on 18 August, 1942, by G. Pereira; SEPFA no. M 14663.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms wide and stiff; tail shorter than head and body; setiforms Ochraceous-Buff; incisive foramen long; vomerine sheath incomplete, or rarely complete; molariform teeth with two counterfolds usually fused.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 22 to 26 mm; maximum width, 0.8 mm. Aristiforms on outer thighs: Gray basally, gradually blackening toward tip but interrupted by Ochraceous-Buff subapical zone; some Ochraceous-Buff to tip; total length, 18 to 20 mm; maximum width, 0.7 mm. Setiforms on middorsal region: Gray basally, gradually blackening toward tip but interrupted by Ochraceous-Buff, subapical zone; total length, 17 to 20 mm; maximum width, 0.06 mm. Setiforms on outer thighs: Gray basally, gradually blackening toward tip but interrupted by Ochraceous-Buff, subapical zone; only a short blackened tip.

Skull.—Large, with elongate rostrum; bullae large and well inflated; jugals with transverse ridge inconspicuous; postorbital process of zygoma small, formed mostly by squamosal; incisive foramen elongated (5.5 × 2.5 mm); vomerine sheath incomplete or, if complete, with maxillary part thin and delicate; posterior palatine foramen at plane of first molars; mesopterygoid fossa extending forward as far as middle parts of second molars.

Teeth.—Upper molariform teeth with two counterfolds; these completely separated in 3 of 16 specimens; two counterfolds coalesced in all three molars in 6 specimens; counterfolds coalesced in only two molars in 3 specimens; counterfolds coalesced in only one molar in 4 specimens. Lower molariform teeth with two counterfolds which are completely separated in 13 of 16 specimens; counterfolds coalesced in only one molar in 2 specimens; counterfolds coalesced in all three molars in one specimen.

Comparisons.—From P. i. gratiosus, bonafidei differs in: Aristiforms wider; tail shorter; molariform teeth with two counterfolds instead of one or two. Differences from P. i. iheringi are given in the account of that subspecies.

Remarks.—Of females with embryos two were captured in April and one in September. The embryos number 2, 1, 2. Young were captured mostly in April, but two were taken in July. Male gonads seemed to be most active in March, April and September. The animals lived in a second growth forest, approaching the climax. The rainfall was more than 1600 mm annually, and the mean annual temperature was 18.5° centigrade.

Specimens examined.—Total number, 18 (MN), from Brazil, Rio de Janeiro, TeresÓpolis, Fazenda BÕa FÉ.

Proechimys iheringi gratiosussubspecies nova

Type locality.—Floresta da Caixa Dagua, Santa Teresa, Espirito Santo, Brazil; altitude 750 meters. Type: Museu Nacional, no. 4024, adult male; collected on 25 May, 1940, by C. Lako; SEPFA no. M 6911.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms narrow; tail of same length as head and body; setiforms Ochraceous-Buff; incisive foramen long; vomerine sheath usually incomplete; upper molariform teeth with one or two counterfolds; lower molariform teeth with two counterfolds, except that m3 usually has only one.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 21 to 27 mm; maximum width, 0.6 mm. Aristiforms on outer thighs: Gray basally, gradually blackening toward middle, and Ochraceous-Buff on distal half; total length, 18 to 21 mm; maximum width, 0.5 mm. Setiforms on middorsal region: Gray basally, gradually blackening toward tip but interrupted by short, Ochraceous-Buff, subapical zone; total length, 18 to 20 mm; maximum width, 0.06 mm. Setiforms on outer thighs: Gray basally, gradually blackening toward middle, and distal part Ochraceous-Buff or with only tip blackened; total length, 14 to 16 mm; maximum width, 0.05 mm.

Skull.—Slender; bullae small but well-inflated; upper edge of jugals deeply concave; transverse ridge of jugals conspicuous; postorbital process of zygoma small, involving only squamosal; incisive foramen elongate (5 x 2.5 mm); vomerine sheath almost always incomplete, and maxillary part lacking or, when present, slender; mesopterygoid fossa extending forward as far as middle of second molars; posterior palatine foramina at plane of front border of first molars or slightly anterior thereto.

Teeth.—Upper molariform teeth with two counterfolds in 10 of 16 specimens and only one in remainder; these folds commonly coalesced; M3 with only one counterfold in 6 specimens, and 2 counterfolds in remainder. Lower molariform teeth with two counterfolds in 6 specimens and in 10 of them m3 has only one counterfold.

Comparisons.—From P. i. panema, gratiosus differs in: Lower molariform teeth with only one counterfold in smaller percentage of specimens; incisive foramen shorter; aristiforms narrower; setiforms Ochraceous-Buff instead of Cinnamon. Differences from iheringi and paratus are given in the accounts of those subspecies.

Remarks.—All the animals were captured in climax forest.

Specimens examined.—Total number, 16 (MN), from Brazil, Espirito Santo, Santa Teresa, Floresta da Caixa Dagua, altitude 750 meters.

Proechimys iheringi panemasubspecies nova

Type locality.—Campinho, Colatina, Espirito Santo, Brazil; altitude 500 meters. Type: Museu Nacional, no. 8288, adult female; collected on 15 July, 1942, by C. Lako.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms moderately wide; tail of approximately same length as head and body; setiforms Cinnamon; incisive foramen moderately long and narrow; vomerine sheath incomplete; upper molariform teeth with two counterfolds, but m3 most frequently with one.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 21 to 23 mm; maximum width, 0.8 mm. Aristiforms on outer thighs: Gray, some gradually blackening toward tip and others with distal part Cinnamon; total length, 17 to 19 mm; maximum width, 0.7 mm. Setiforms on middorsal region: Gray, gradually blackening toward tip, but interrupted by Cinnamon, subapical zone; total length, 18 to 20 mm; maximum width, 0.06 mm. Setiforms on outer thighs: Gray, gradually blackening toward middle, and Cinnamon on all of distal parts or with tip blackish; total length, 13 to 15 mm; maximum width, 0.09 mm.

Skull.—Strong, with jugals dorso-ventrally wide; interorbital region and cranium wide; bullae well inflated; transverse ridge of jugals not well-developed; postorbital process of zygoma small and formed only of squamosal; incisive foramen 4.7 × 2.2 mm; vomerine sheath always incomplete, with maxillary part reduced to small process; mesopterygoid fossa extending forward as far as middle of second molars or only slightly short thereof; posterior palatine foramina at plane of front of first molars.

Teeth.—All upper molariform teeth with two counterfolds in 4 specimens; one having only one counterfold in M3; 3 with counterfolds coalesced in one or two molars. Lower molariform teeth with two counterfolds in one specimen, these counterfolds not coalesced; m3 with one counterfold in 4 specimens and with two in one specimen.

Comparisons.—Differences from P. denigratus and P. i. paratus are given in the accounts of those animals.

Specimens examined.—Total number, 5 (MN), from Brazil, Espirito Santo, Colatina, Campinho; altitude 500 meters.

Proechimys iheringi denigratussubspecies nova

Type locality.—Mata do RibeirÃo da Fortuna, 40 kilometers west of Ilheus, Itabuna, Bahia, Brazil. Type: Museu Nacional, no. 8500, adult male; collected 16 March, 1945.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms wide and stiff; tail longer than head and body; setiforms near (15"a) Cinnamon; incisive foramen long and narrow; vomerine sheath complete; premolars with two counterfolds, upper molars with one or two, and lower molars with only one.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 20 to 22 mm; maximum width, 1.1 mm. Aristiforms on outer thighs: Gray basally, gradually blackening toward tip or with distal part near (15"a) Cinnamon; total length, 14 to 16 mm; maximum width, 0.5 mm. Setiforms on middorsal region: Gray basally, gradually blackening toward tip but interrupted by near (15"a) Cinnamon, subapical zone 4 mm wide; total length, 18 to 20 mm; maximum width, 0.05 mm. Setiforms on outer thighs: Gray basally, gradually blackening toward tip but interrupted by wide, near (15"a) Cinnamon, subapical zone.

Skull.—Slender; nasals short; bullae large and well-inflated; jugals with conspicuous transverse ridge; postorbital process of zygoma conspicuous, spiniform and formed almost exclusively by jugal; incisive foramen elongated and narrow (5 × 1.8 mm); vomerine sheath complete and formed almost exclusively by premaxillae; maxillary part of this sheath short and in most specimens the two parts of sheath completed by vomer itself; mesopterygoid fossa extending forward as far as middle of second molars and in some skulls as far as anterior border of second molars; posterior palatine foramina at anterior plane of first molars.

Teeth.—Upper molariform teeth: P4 always with two counterfolds; M1 with two counterfolds in 65 per cent of specimens but anterior counterfold poorly developed; rest of specimens with only one counterfold in M1; M2 with two counterfolds in 50 per cent of specimens and only one in remainder; M3 with two counterfolds in only 17 per cent of specimens, and remainder with only one. Lower molariform teeth: p4 always with two counterfolds; molars always with only one counterfold.

Comparisons.—From P. i. panema, denigratus differs in: Each lower molar with only one, instead of with more than one, counterfold; incisive foramen longer and narrower; vomerine sheath complete instead of incomplete; aristiforms conspicuously wider; tail longer.

Remarks.—One female (SEPFA no. M 17060) captured on 9 January, 1944, gave birth to two females on 26 January, 1944. Each of these young measured 177 mm in total length and weighed 27.8 g. On 4 March, 1944, their measurements were: head and body, 120,120; tail, 120,130; hind foot, 32,33; ear, 21,22; skull:—total length, 36.0,35.0; condyloincisive length, 29.0,29.1; zygomatic breadth, 19.1,18.5; length of nasals, 12.5,11.6; interorbital constriction, 9.3,8.8; cranial breadth, 16.4,16.9; palatilar length, 11.5,10.5; crown length of P4 and M1, 4.3,4.3 mm.

The forest where the animals were captured has a high percentage of deciduous trees in spite of the heavy rainfall in this region. All of the animals were trapped near water. Young were captured from January to May. Most animals have a conspicuous Cinnamon patch on the nuchal region.

Specimens examined.—Total number, 34 (SEPFA 33, MN 1), from Brazil, Bahia, Itabuna, Mata do RibeirÃo da Fortuna.

Proechimys iheringi paratussubspecies nova

Type locality.—Floresta da Capela de SÃo Braz, Santa Teresa, Espirito Santo, Brazil; altitude 630 meters. Type: Museu Nacional, no. 4012, adult female; collected on 24 September, 1940, by Dr. H. W. Laemmert; SEPFA no. M 8447.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms wide and stiff; tail 96 per cent of head and body; color on setiform Cinnamon-Buff; incisive foramen short and moderately wide; vomerine sheath complete; all molariform teeth with two counterfolds.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 24 to 26 mm; maximum width, 1.3 mm. Aristiforms on outer thighs: Gray basally, gradually blackening toward middle, and distal parts near (15"c) Pinkish Cinnamon; total length, 18 to 20 mm; maximum width 0.8 mm. Setiforms on middorsal region: Gray basally, gradually blackening toward tip but interrupted by Cinnamon-Buff, subapical zone; total length 14 to 16 mm; maximum width, 0.06 mm.

Skull.—Slender; bullae large and well-inflated; jugals with conspicuous, transverse ridge; postorbital process of zygoma moderately developed and involving only squamosal; incisive foramen short and narrow (4.1 × 2.1 mm); vomerine sheath complete, with maxillary part short and thick; mesopterygoid fossa extending forward as far as posterior parts of second molars; posterior palatine foramina at plane of premolars.

Fig. 128. Fig. 128.Map showing the geographic ranges of the subspecies of Proechimys iheringi.

Teeth.—Upper and lower molariform teeth with two counterfolds. Counterfolds coalesced in P4 and M1 of one specimen.

Comparisons.—From P. i. gratiosus and P. i. panema, paratus differs in: all molariform teeth with two, instead of some with fewer, counterfolds; vomerine sheath complete and thick instead of usually incomplete; incisive foramen shorter and narrower; aristiforms conspicuously wider; setiforms Cinnamon-Buff instead of Ochraceous-Buff and Cinnamon, respectively. Tail 96 per cent of head and body in paratus instead of 100 per cent as in panema.

Remarks.—The animals were captured in climax forest.

Specimens examined.—Total number, 3 (MN), from Brazil, Espirito Santo, Santa Teresa, Floresta da Capela de SÃo Braz; altitude 630 meters.

Proechimys setosus (Desmarest)

General characters.—Size medium; tail approximately same length as head and body; aristiforms moderately wide; feet rather large; ears of medium size; color on upper parts and sides sepia gradually changing to Ochraceous-Tawny; few differentiated, light-colored aristiforms present on outer thighs and rump; under surface of body and inner sides of legs white; tail with white tip and conspicuous, white pencil; feet white dorsally; skull short and smooth, somewhat flattened in interorbital region; jugals narrow dorso-ventrally; incisive foramen moderately long and notably narrow; vomerine sheath complete and slender; postorbital process of zygoma spinelike and involving mostly jugal; premolars usually with two counterfolds; molars with only one counterfold, rarely two in M1 or in M3.

Remarks.—The specimens available are undoubtedly faded and, therefore, the colors mentioned above for the upper parts and sides may not correspond to the colors of unfaded pelages. Desmarest (1817:59) describes the color of setosus as similar to that of the "Echimys de Cayenne" (Proechimys guyannensis) but being more "rousse." Is. Geoffroy Saint-Hilaire (1840:52) describes the same animal as being "d'un brun roussÂtre" on the upper parts.

The Proechimys from Lagoa Santa, Minas Gerais, "Echimys" elegans Lund, is certainly related to P. iheringi as well as to P. albispinus. From P. iheringi, elegans differs in having a smaller skull with shorter rostrum, narrower incisive foramen, and orthodont incisors. On the other hand the restricted distribution of the aristiforms in the pelage and the white, penicillate tail are points of resemblance to iheringi. From P. albispinus, elegans differs in having a less spinous pelage and longer tail with white pencil instead of a brown pencil. The skulls, however, are similar, except for the fact that elegans does not have proodont incisors as albispinus sometimes does. Thomas (1921:141) states, after describing the skull of the type of setosus, that "Specimens corresponding to this animal have been obtained at Lagoa Santa, Minas, by Lund and others, and at Bahia." Thomas, however, would not have referred to specimens from "Bahia" as being comparable to elegans had they not been different from albispinus which he discussed in the same paper. Also, he would not have confused "specimens comparable to elegans" with a subspecies of P. iheringi (P. i. denigratus, from southern Bahia) which has opisthodont instead of orthodont incisors. Since French collectors sent material to Europe at the beginning of the 19th century from (southern?) Bahia, possibly setosus came from there.

In the collection of the American Museum of Natural History there is one specimen (AMNH no. 16140) of Proechimys, included in the so-called Maximilian Collection. The characters of this specimen agree closely with those of the specimens from Lagoa Santa. The locality of capture of specimen no. 16140 is unknown, but it is reasonable to assume that Prince Maximilian zu Wied obtained it somewhere along his route of travel through southeastern Bahia. Wied (1826:445) mentions "L[oncheres]. myosuros Licht." as "am Parahyba, am Peruhype und Belmonte," which greatly increases the possibility of its having come from southern Bahia. The close similarity to elegans of Wied's specimen indicates that the locality of capture possibly was in the region of the less humid, low escarpments of southern Bahia.

My conclusion is that Wied's specimen corresponds closely to setosus and, tentatively, I identify it as such. "Echimys elegans," due to the relationships mentioned above is here considered to be a subspecies of setosus.

Among the species described in earlier times, and whose identity was never ascertained, "Echinomys" fuliginosus Wagner seems to be synonymous with setosus. Wagner describes the animal as having a tail "apicis versus pilis albidis vestita" and the figure of the cheekteeth (1844, pl. 239 D) shows a typical trilaminate condition which occurs commonly in elegans. Moreover, the tail of fuliginosus is only 9 per cent shorter than the head and body and the aristiforms of this subspecies are moderately wide.

Proechimys setosus setosus (Desmarest)

Echimys setosus Desmarest (Geoffroy's MS), 1817, Nouv. Dict. Hist. Nat. nouv. ed., 10:59 (orig. descr.); Is. Geoffroy Saint-Hilaire, 1838, Comptes Rendus Acad. Sci., Paris, 6(26):886; Is. Geoffroy Saint-Hilaire, 1840, Mag. Zool., Paris, (ser. 2, annÉe 2):12, 33, 52; Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20):257, 261.

Echimys cayennensis Pictet, 1841, MÉm. Soc. phys. Hist. Nat., GenÈve, 9:145; Waterhouse, 1848, Nat. Hist. Mammalia, 2:334.

Echinomys fuliginosus Wagner, 1843, Schreber's SÄugethiere, suppl. 3:343; Wagner, 1844, Schreber's SÄugethiere, suppl. 4, pl. 39 D.

Proechimys setosus Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20):264; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):141; Tate, 1935, Bull.

Amer. Mus. Nat. Hist., 68(5):401; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.

Proechimys fuliginosus Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5): 400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:119.

Type locality.—Unknown; see remarks under P. setosus. Type: Museum d'Histoire Naturelle, Paris, no. A. 7787 (Thomas, 1921:141), "very imperfect."

Diagnosis.—Aristiforms wide; P4 and M1 with two counterfolds; p4 with two counterfolds, one anterior to main fold.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 18 to 20 mm; maximum width, 0.8 mm. Aristiforms on outer thighs: Color much faded; total length, 15 to 17 mm; maximum width, 0.3 mm. Setiforms on middorsal region: Color faded; total length, 16 to 18 mm; maximum width, 0.04 mm. Setiforms on outer thighs: Color faded; total length, 10 to 13 mm; maximum width, 0.03 mm.

Figs. 129-132.

Figs. 129-132.Proechimys setosus elegans, sex ?, UZM no. L 104, Lagoa Santa. × 1.

Skull.—Short; rostrum short and stout; length of nasals approximately 15 mm (broken); bullae roundish, smooth and well-inflated; jugals dorso-ventrally narrow (3.1 mm) with strong transverse ridge; postorbital process of zygoma spiniform, slender and involving mostly jugal; incisive foramen narrow (3.8 x 1.7 mm) and narrowest in posterior part; vomerine sheath complete; posterior palatine foramina obsolete; mesopterygoid fossa extending forward as far as middle of second molars.

Teeth.—Incisors orthodont. P4 with two counterfolds; M1 with two counterfolds but anterior one notably small; M2 and M3 with only one counterfold each. In lower jaw: p4 with two counterfolds, one anterior to main fold; molars with only one counterfold.

Comparisons.—From P. s. elegans, setosus differs in: M1 with two counterfolds as opposed to only one; M3 with one counterfold instead of sometimes with two counterfolds; p4 with one counterfold anterior to main fold and another posterior, instead of both counterfolds posterior.

Remarks.—The measurements above were taken from the Maximilian specimen mentioned above. Measurements of the type were given by Desmarest as: head and body, 5-1/2 inches, tail about 6-1/2 inches. Is. Geoffroy Saint-Hilaire (1838:886) corrects these measurements to: head and body 195 mm; tail (part missing), 170 mm.

Proechimys setosus elegans (Lund)

E[chimys]. elegans Lund, 1841, Kong. Danske Videnskab. Selsk. natur-vidensk. math. Afhandl., KjÖbenhavn, 8:99 (orig. descr.).

Loncheres elegans Lund, 1841, Kong. Danske Videnskab. Selsk, natur-vidensk. math. Afhandl., KjÖbenhavn, 8:245, 266, 294; Wagner, 1843, Wiegman's Archiv f. Naturg., Berlin, 2 (Jahrg. 9):47.

Echimys cayennensis Waterhouse, 1848, Nat. Hist., Mammalia, 2:337.

Echinomys cajennensis Winge, 1888, Jordfundne og nulevende Gnavere (Rodentia), E Museo Lundii, KjÖbenhavn, 1(3):71, pl. 6, figs. 5-6, pl. 7, fig. 1.

Proechimys setosus Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):141.

Proechimys elegans Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):400; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:119.

Type locality.—Lagoa Santa, Nova Lima, Minas Gerais, Brazil. Type: Syntypes in Universitets Zoologiske Museum, KjÖbenhavn; collected by P. W. Lund.

Range.—Known only from the type locality.

Diagnosis.—Aristiforms wide; P4 usually with two counterfolds; M3 sometimes with two counterfolds; p4 with two counterfolds anterior to main fold.

Pelage.Aristiforms on middorsal region: Gray basally, gradually blackening toward tip; total length, 18 to 20 mm; maximum width, 0.7 mm. Aristiforms on outer thighs: Gray basally, gradually blackening toward tip which is Cinnamon; total length, 15 to 17 mm; maximum width, 0.3 mm. Setiforms on middorsal region: Whitish basally, gradually blackening toward tip, but interrupted by Cinnamon, subapical zone; total length, 17 to 19 mm; maximum width, 0.04 mm. Setiforms on outer thighs: Whitish basally, gradually blackening toward tip but interrupted by near (15"a) Cinnamon, subapical zone; total length, 10 to 12 mm; maximum width, 0.03 mm.

Skull.—Short; rostrum short but not stout; length of nasals 17 mm; bullae large, smooth, and well-inflated; jugals with conspicuous, transverse ridge; postorbital process of zygoma long, spiniform and constructed entirely of jugal; incisive foramen narrow (4 × 1.7 mm); vomerine sheath complete and slender; posterior palatine foramina obsolete; mesopterygoid fossa extending anteriorly as far as middle parts of second molars.

Teeth.—Incisors orthodont. P4 usually with two counterfolds, rarely with three; upper molars with only one counterfold, but M3 sometimes with two, posterior one being vestigial. Lower molariform teeth: p4 with two counterfolds, both being anterior to main fold; molars with only one counterfold.

Comparisons.—Differences from P. s. setosus are given in the account of that subspecies.

Remarks.—According to Lund, these animals are found in the vicinity of small pools, swim well in spite of not having webbed toes, at night go after food and climb the corn stalks, and have their nests in the grass at the margins of the pools.

Specimens examined.—Total number, 2 (UZM), from Brazil, Minas Gerais, Nova Lima, Lagoa Santa.

Proechimys albispinus(Is. Geoffroy)

General characters.—Size small; tail of same length as head and body or slightly less; feet small; ears of medium size; color of upper parts Ochraceous-Tawny gradually changing to Ochraceous-Buff on sides; differentiated, light-colored aristiforms on back, sides, rump and at base of tail; clavate aristiforms on back with Ochraceous-Tawny or Ochraceous-Buff, subapical zone; underparts of body and inner sides of legs white; tail blackish above, white below, with no white tip; hands and feet white on dorsal parts and some specimens darker on outer margins of feet; skull short and smooth, somewhat flattened in frontal region; jugal dorso-ventrally wide and with moderately conspicuous transverse ridge; postorbital process of zygoma well developed and involving both jugal and squamosal; bullae large and smooth; incisive foramen short and narrow; vomerine sheath incomplete or complete; molariform teeth with only one counterfold; incisors orthodont or proodont.

Remarks.—A good series from Macaco Seco, AndaraÍ, Bahia, agrees closely with the form first described (albispinus) from the Island Madre de Deus, in Todos os Santos Bay, Bahia. Compared with topotypes of P. albispinus sertonius, the animal from Macaco Seco in general color is more Ochraceous-Tawny and has a narrower skull with orthodont incisors. Specimens from Bonfim, northeastern Bahia, on the other hand, agree with Thomas' albispinus sertonius, from LamarÃo, being browner and having broader skulls than P. a. albispinus and having proodont, instead of orthodont, incisors. The range of each of the two subspecies is, therefore, fairly extensive. The insular form extends to the less rainy, continental area and the form from LamarÃo ranges northward (NNW) in the same type of highly deciduous forest, the "caatinga."

The species albispinus is certainly the most specialized form of the entire genus for drier habitats. In addition to the general adaptations described above, it is noteworthy for having both lanceolate and clavate aristiforms. The latter type has a wide basal part and an abruptly narrowed, distal part. The same development is seen in the genus Echimys, where highly spinous forms, like Echimys paleacea (Lichtenstein), show the same two types of aristiforms.

Figs. 133-140.

Figs. 133, 135.Proechimys albispinus albispinus, male, CNHM no. 20409, Macaco Seco. × 1.

Figs. 134, 136.Proechimys albispinus sertonius, male, MN no. 6454, Bonfim. × 1

Figs. 137, 138.Proechimys albispinus albispinus, male, CNHM no. 20409, Macaco Seco. × 1.

Figs. 139, 140.Proechimys albispinus sertonius, male, MN no. 6454, Bonfim. × 1.

Proechimys albispinus albispinus(Is. Geoffroy)

Echimys albispinus Is. Geoffroy Saint-Hilaire, 25 June 1838, Comptes Rendus Acad. Sci., Paris, 6(26):886; Is. Geoffroy, August, 1838, Ann. Sci. Nat., Paris, 10 (ser. 2):125; Is. Geoffroy, 1840, Mag. Zool., Paris (ser. 2, annÉe 2, livr. 13):33, 53, pl. 26, pl. 29 (figs. 1, 2, 3); Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20):261.

Echinomys fuliginosus Wagner, 1843, Schreber's SÄugethiere, suppl., 3:343.

Echimys albispinosus Waterhouse, Nat. Hist., Mammalia, 2:341.

Proechimys albispinus Allen, 1899, Bull. Amer. Mus. Nat. Hist., 12(20):264; Thomas, 1911, Ann. Mag. Nat. Hist., 8 (ser. 8):252.

Proechimys albispinus albispinus Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):141; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):401; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.

Type locality.—Ilha Madre de Deus, Itaparica (near Salvador), Bahia, Brazil. Type: Museum d'Histoire Naturelle, Paris, no. A 7669, "skull ... practically perfect" (Thomas, 1921:142).

Range.—Island Madre de Deus, Macaco Seco, AndaraÍ and probably islands of the bay of Todos os Santos and valley of the ParaguassÚ River.

Diagnosis.—Aristiforms wide; color on setiforms Ochraceous-Tawny on upper parts and sides; incisors orthodont; molariform teeth with one counterfold, p4 rarely with two.

Pelage.Aristiforms on middorsal region: Lanceolate aristiforms, with basal part whitish, gradually blackening toward tip; total length, 25 to 28 mm; maximum width, 1.2 mm; clavate aristiforms with base whitish, gradually blackening toward tip but interrupted by Ochraceous-Tawny, subapical zone. Aristiforms on outer thighs: Whitish on basal half, gradually blackening toward tip; total length, 24 to 26 mm; maximum width, 0.9 mm. Some are whitish basally, gradually blackening toward distal part but distal fourth or fifth near (15'j) Ochraceous-Tawny. Setiforms on middorsal region: Whitish basally, gradually blackening toward tip but interrupted by Ochraceous-Tawny, subapical zone; total length, 20 to 23 mm; maximum width, 0.1 mm. Some setiforms almost completely whitish. Setiforms on outer thighs: Whitish basally, gradually blackening toward tip but interrupted by near (15'j) Ochraceous-Tawny, subapical zone; total length, 18 to 20 mm; maximum width, 0.06 mm.

Skull.—Narrow; bullae small and smooth; jugals dorso-ventrally wide with conspicuous transverse ridge; postorbital process of zygoma well-developed and formed by jugal and squamosal; posterior palatine foramina obsolete; incisive foramen narrow and short; vomerine sheath complete or incomplete but premaxillary part at a level lower than that of maxillary part (when skull is viewed from ventral face); mesopterygoid fossa extending forward as far as anterior borders of second molars.

Teeth.—Incisors orthodont; molariform teeth with only one counterfold, except that p4 rarely has two counterfolds.

Comparisons.—From P. a. sertonius, albispinus differs in: sides of body darker; incisors orthodont as opposed to proodont; p4 rarely with two counterfolds instead of always with one counterfold.

Remarks.—The localities where P. a. albispinus has been collected have a forest climax with a moderate percentage of deciduous trees.

Specimens examined.—Total number, 19 (18 CMNH, 1 MCZ), from Brazil, Bahia, AndaraÍ, Macaco Seco.

Proechimys albispinus sertoniusThomas

Proechimys albispinus sertonius Thomas, July, 1921, Ann. Mag. Nat. Hist., 8 (ser. 9):142 (orig. descr.); Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):401; Ellerman, 1940, The families and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:122.

Type locality.—LamarÃo ("about 70 miles north of Bahia City"), Ituiutaba, Bahia, Brazil; altitude 300 meters. Type: British Museum (Nat. Hist.), no. 3.9.5.86, adult male; collected on 16 June, 1903, by Alphonse Robert; original number, 1508.

Range.—Known from the type locality and Bonfim; probably occupies valleys of Jacuipe and the ItapicurÚ rivers and littoral between them.

Diagnosis.—Aristiforms wide; color of setiforms Ochraceous-Tawny on back, grading to Ochraceous-Buff on sides; incisors proodont; no molariform tooth with more than one counterfold.

Pelage.Aristiforms on middorsal region: Lanceolate aristiforms whitish basally, gradually blackening toward tip; total length, 23 to 27 mm; maximum width, 1.3 mm. Clavate aristiforms, and some lanceolate ones, whitish basally, gradually blackening toward tip but interrupted by Ochraceous-Tawny, subapical zone. Some clavate aristiforms without subapical zone but blackened in distal part; total length, 23 to 24 mm; maximum width, 0.7 mm. Aristiforms on outer thighs: Whitish basally, gradually blackening toward tip but interrupted by Light Ochraceous-Buff, subapical zone; tip slightly darker; some whitish basally, grayish in middle and light yellowish toward tip; total length, 20 to 22 mm; maximum width, 0.9 mm. Setiforms on middorsal region: Whitish basal part succeeded by grayish, then by long, light, yellowish band, which becomes Light Ochraceous-Buff, and blackish tip; total length, 26 to 29 mm; maximum width, 0.15 mm. Setiforms on outer thighs: Whitish basal part succeeded by grayish, then Light Ochraceous-Buff, subapical zone and blackish tip; total length, 18 to 20 mm; maximum width, 0.13 mm.

Skull.—Broad; bullae small and smooth; jugals dorso-ventrally "wide," with conspicuous transverse ridge; postorbital process of zygoma well-developed and formed by both jugal and squamosal; incisive foramen narrow and short; vomerine sheath incomplete or complete but premaxillary part on a lower level than maxillary part (when skull is viewed from ventral face); mesopterygoid fossa extending forward as far as anterior faces of second molars.

Teeth.—Incisors proodont; molariform teeth with only one counterfold.

Comparisons.—Differences from P. a. albispinus are given in the account of that subspecies.

Remarks.—Localities where samples were collected are typical "caatinga" forest, a climax of mainly deciduous trees; cacti are also common in the region.

Specimens examined.—Total number, 10, from Brazil, Bahia, as follows: Ituiutaba, LamarÃo, 4 (1 DZ, 1 CMNH, 1 MCZ, 1 USNM); Bonfim, Bonfim, 6 (5 DZ, 1 MN).


                                                                                                                                                                                                                                                                                                           

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