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diff --git a/42720-0.txt b/42720-0.txt new file mode 100644 index 0000000..e8b1a26 --- /dev/null +++ b/42720-0.txt @@ -0,0 +1,4609 @@ +*** START OF THE PROJECT GUTENBERG EBOOK 42720 *** + + Speciation in the Brazilian Spiny Rats + (Genus Proechimys, Family Echimyidae) + + BY + JOÃO MOOJEN + + + University of Kansas Publications + Museum of Natural History + + Volume 1, No. 19, pp. 301-406, 140 figures in text + December 10, 1948 + + + University of Kansas + LAWRENCE + 1948 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, A. Byron Leonard, + Edward H. Taylor + + Volume 1, No. 19, pp. 301-406, 1 plate, 140 figures in text + + Published December 10, 1948 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED BY + FERD VOILAND, JR., STATE PRINTER + TOPEKA, KANSAS + 1948 + + 22-3343 + + + + + Speciation in the Brazilian Spiny Rats + (Genus Proechimys, Family Echimyidae) + + By + JOÃO MOOJEN + + + + +CONTENTS + + + PAGE + INTRODUCTION 305 + + METHODS AND TERMINOLOGY 305 + + ACKNOWLEDGMENTS 308 + + PALEONTOLOGY 308 + + SPECIATION 311 + Subgeneric variation 312 + Specific variation in the subgenus _Proechimys_ 314 + Subspecific variation in the subgenus _Proechimys_ 317 + Specific variation in the subgenus _Trinomys_ 320 + Subspecific variation in the subgenus _Trinomys_ 322 + + TAXONOMIC CHARACTERS 323 + Size and proportions of external parts 323 + Pelage 324 + Skull 326 + Incisive foramen 326 + Teeth 327 + + HABITS 330 + + CHANGES WITH AGE 331 + + Genus PROECHIMYS 333 + + ARTIFICIAL KEY TO SUBGENERA AND SPECIES 334 + + Subgenus PROECHIMYS 338 + _Proechimys goeldii_ 338 + _Proechimys goeldii steerei_ 338 + _Proechimys goeldii goeldii_ 340 + _Proechimys semispinosus_ 342 + _Proechimys semispinosus liminalis_ 343 + _Proechimys semispinosus amphichoricus_ 344 + _Proechimys semispinosus kermiti_ 345 + _Proechimys longicaudatus_ 346 + _Proechimys longicaudatus brevicauda_ 349 + _Proechimys longicaudatus boimensis_ 350 + _Proechimys longicaudatus longicaudatus_ 351 + _Proechimys longicaudatus leucomystax_ 352 + _Proechimys longicaudatus roberti_ 353 + _Proechimys guyannensis_ 355 + _Proechimys guyannensis villicauda_ 355 + _Proechimys guyannensis ribeiroi_ 361 + _Proechimys guyannensis hyleae_ 361 + _Proechimys guyannensis nesiotes_ 363 + _Proechimys guyannensis leioprimna_ 364 + _Proechimys guyannensis oris_ 365 + _Proechimys guyannensis arescens_ 366 + _Proechimys guyannensis riparum_ 367 + _Proechimys guyannensis arabupu_ 369 + + Subgenus TRINOMYS 369 + _Proechimys dimidiatus_ 371 + _Proechimys iheringi_ 373 + _Proechimys iheringi iheringi_ 378 + _Proechimys iheringi bonafidei_ 378 + _Proechimys iheringi gratiosus_ 379 + _Proechimys iheringi panema_ 380 + _Proechimys iheringi denigratus_ 381 + _Proechimys iheringi paratus_ 382 + _Proechimys setosus_ 384 + _Proechimys setosus setosus_ 385 + _Proechimys setosus elegans_ 387 + _Proechimys albispinus_ 388 + _Proechimys albispinus albispinus_ 390 + _Proechimys albispinus sertonius_ 391 + + INCERTA SEDIS 392 + _Proechimys myosuros_ 392 + + CONCLUSIONS 393 + + TABLE OF MEASUREMENTS 395 + + LITERATURE CITED 400 + + +[Illustration: FIG. 1. _Proechimys dimidiatus_ (Günther). Live female on +left and male on right. ×1/2. From Tingua, Nova Iguassú, Rio de Janeiro, +Brazil. Photographed in spring (August or September) of 1942 by +author.] + + + + +INTRODUCTION + + +The spiny-rats included in the genus _Proechimys_ are common in almost +every forest of South America above the Tropic of Capricorn, and in +Central America northward to approximately 12° N, in Nicaragua. In size +and proportions they are similar to the brown rat _Rattus norvegicus_ +but actually they belong to a very different suborder of rodents--the +Hystricomorpha. The hystricomorphs are represented in South America by a +large variety of animals, of which capybaras, agoutis and cavies are +common representatives. + +The pelage of the spiny-rats has a large number of flattened, spinelike +hairs, especially on the back. The color ranges through different tints +and shades of reddish-brown more or less evenly distributed on the upper +parts; the underparts are usually pure white, sharply contrasting with +the brown color above. The tail is bicolored, brown above and white +below. + +The spiny-rats live in forests of different types, generally in the +proximity of water. Shelter is usually procured under boulders, stumps +or masses of roots. The reproductive rate is low; on the average, there +are only two young per litter and only two litters per year. + +Sixty-odd names have been given to species and subspecies of +_Proechimys_ in the last hundred and fifty years and no serious revision +of the taxonomy of the genus was undertaken in the last century. The +purpose of the present work is to provide means of understanding species +and subspecies within the genus and to describe the different kinds +known to occur within the confines of Brazil. + + + + +METHODS AND TERMINOLOGY + + + PELAGE.--It was found advisable to use a standardized nomenclature + for hairs. The names here proposed are a choice of those used in + the literature, with the suffix "_form_" as an element of + uniformity. I feel that it would be advantageous if everyone + adopted a similar universal system in mammalogy. + + The names listed below are used as nouns and are considered as + English versions which could easily be adapted to different + languages. These names may be complemented with adjectives as + needed. Examples are lanceolate aristiforms, spinous aristiforms, + and woolly setiforms. + + _Aristiforms:_ The most conspicuously developed hairs in a + three-layered pelage or the corresponding hairs in a simpler + pelage. Names previously used for these hairs are: guard hair, + leithaar and jarre. + + _Setiforms:_ Common to all species and most numerous throughout the + pelage; second in conspicuousness, being the dominant hairs in the + middle layer. Synonyms are: over hairs, grannenhaare and soies. + + _Villiforms:_ The smallest hairs in the three-layered pelage. + Synonyms are: underfur, wollhaar and duvet. + + _Vibrissiforms:_ The vibrissae proper, or any typically sensory + hair. + + TEETH.--The tritubercular nomenclature was abandoned because of + overwhelming difficulties; more research on the Hystricomorpha is + certainly needed before the tritubercular nomenclature can be + applied with confidence. The following names are used for features + of the molariform teeth: + + _Main fold:_ The inner or lingual fold in the upper molariform + teeth and outer or labial fold in the lower molariform teeth. + + _Counterfold:_ Any outer or labial fold in the upper or any inner + or lingual fold in the lower molariform teeth. + + For incisors Thomas (1921:141) is followed: _opisthodont_, + _orthodont_ and _proodont_ depending on the angle between the + exposed part of incisors and the ventral surface of the rostrum. + + The capital letters P and M designate premolars and molars, + respectively, of the upper jaws; lower case letters p and m + designate corresponding teeth in the lower jaws. + + MEASUREMENTS.--Measurements of skins were used only when provided + by the collector. The length of the hind-foot is intended to be + always _cum unguis_, but in a few instances it is impossible to be + sure whether the collector included the nail. Length of tail was + used only when the tail seemed not to be mutilated. Ear + measurements taken by collectors are scarce. In spite of the + apparent usefulness of length of ear, it was found to be + inadvisable to take the measurement on the dry skins. + + The following measurements of the skull are used in the tables: + + _Greatest length:_ From the anteriormost part of the nasals to the + posteriormost part of the supraoccipital. + + _Condylo-incisive length:_ From the anterior face of one incisor, + at the alveolus, to the posteriormost part of the exoccipital + condyle of the same side. + + _Zygomatic breadth:_ Maximum distance across zygomata in a plane + perpendicular to longitudinal axis of the skull. + + _Length of nasals:_ Maximum length of one or both, whichever is + the greater. + + _Interorbital constriction:_ Least width between the orbits on top + of the skull. + + _Palatilar length:_ From the posterior face of an incisor, at the + alveolus, to the nearest part of the posterior edge of the + palatine bone. + + _Crown length of cheekteeth:_ From the anterior border of P4 to + the posterior border of M3. + + In the accounts of species, measurements of aristiforms and + setiforms are used. The hairs measured were taken from the + middorsal region and outer thighs, and the measurements are means. + + All specimens of which measurements are here recorded, as for + example in the tables, are fully adult; each specimen shows some + wear on each of the four upper molariform teeth unless otherwise + indicated. + + Capitalized color terms are after Ridgway "Color Standards and + Color Nomenclature," Washington, D. C., U. S. A., 1912. One + setiform was taken from the animal and placed over the rectangles + in Ridgway's charts and the examination made under a microscope + with low (×7) magnification and natural light. This method was + found to give the most satisfactory results. + +The following abbreviations are used for names of institutions: + + AMNH--American Museum of Natural History. + CNHM--Chicago Natural History Museum. + DZ--Departamento de Zoologia da Secretaria de Agricultura, São + Paulo, Brazil. + MCZ--Museum of Comparative Zoology at Harvard College. + MN--Museu Nacional, Brazil. + MZ--Museum of Zoology, University of Michigan. + SEPFA--Serviço de Estudos e Pesquisas sobre a Febre Amarela, Brazil. + USNM--United States National Museum. + UZM--Universitets Zoologiske Museum, Copenhagen. + + + + +ACKNOWLEDGMENTS + + + Approximately two thousand skins and skulls were assembled at the + Museum of Natural History, University of Kansas, through the + coöperation of the authorities in the various institutions of North + America, Brazil and Denmark, as listed immediately above. This + comprehensive material was used to obtain a more complete + understanding of the group, and for the loan of these specimens I + am extremely grateful to the authorities of each of the + institutions. + + First of all I acknowledge the encouragement given me in the + _Proechimys_ project by Heloisa Alberto Torres, Director of the + Museu Nacional, Rio de Janeiro. I extend my thanks also to Stephen + D. Durrant, of the University of Utah, for helpful corrections in + the preparation of the manuscript; to Mrs. Virginia Cassell Unruh, + for the preparation of the drawings of the skulls; to Miss Alice + M. Bruce for assistance in drawing the maps; and to my daughter, + Julieta, for help in assembling data and for typing. + + Dr. Remington Kellogg, Curator of Mammals in the United States + National Museum, and the late Dr. Wilfred H. Osgood, formerly + Curator Emeritus of the Department of Zoology in the Chicago + Natural History Museum, generously permitted me to use their + private lists of South American mammals. These lists contain much + unpublished data, as for example, proof, in Kellogg's list, that + _Proechimys guyannensis_ (E. Geoffroy Saint-Hilaire, 1803) + antedates _P. cayennensis_ (Desmarest, 1817). I register here my + gratitude to both these zoölogists and acknowledge other critical + assistance from Dr. Kellogg. + + The John Simon Guggenheim Memorial Foundation awarded me a + fellowship for which I am deeply grateful. This expression of the + Foundation's interest in education and good neighborliness made + possible the completion of the present paper. + + Finally I desire to express my deepest gratitude to Professor E. + Raymond Hall, Director of the Museum of Natural History and + Chairman of the Department of Zoology at the University of Kansas + whose untiring aid and guidance has enabled me to terminate this + study. + + + + +PALEONTOLOGY + + +The only known, significant, fossil _Proechimys_ comes from deposits in +the limestone caves of Lagoa Santa, Minas Gerais, Brazil. These +deposits, of Late Pleistocene or Recent age, were extensively studied by +P. W. Lund and the results published in a series of French and Danish +papers. F. Ameghino (1934:110) studied another fauna from a deposit of +similar age in the cave of Iporanga, São Paulo, Brazil. _Proechimys_ is +recorded in his account under the inclusive specific name _fuliginosus_. + +The molariform teeth of the fossil described by Lund (1841:pl. 21, fig. +14) shows its close relationship to the living form _P. s._ _elegans_ +(Lund) which still inhabits the same region. It belongs in the more +specialized subgenus _Trinomys_ which seems to have been derived from +_Proechimys_. _Trinomys_ has the main fold in the molars always greatly +developed and the fold tends to set apart one lamina in the occlusal +surface. The Lagoa Santa fossil, like some specimens of the living +subspecies, has a small main fold in P4. However, the main fold is large +in all upper molars and in the lower molariform teeth which are notably +specialized in the extreme reduction of the number of counterfolds to +only one. + +One hypothesis concerning the evolution of the genus is that a more +primitive group of _Proechimys_ lived in all of the Central Plateau of +Brazil in the Pleistocene Time. The climatic conditions at that time +might have been such as to support large forests but, since the +Pleistocene, these climatic conditions may have changed from humid to +the present drier conditions, which support the dominant, savanna, +floral climax. Actually the extinct fauna from the caves includes +animals which have disappeared from the area and now live only in more +humid areas, as for example _Myocastor_, which has shifted to the +lowlands to the west and south. + +Possibly climatic changes were responsible for the faunal shift from the +region that is now a plateau in Central Brazil. This climatic change may +have resulted from the gradual uplift of the eastern part of the +continent. This uplift prevents part of the trade winds which come from +the east from carrying the same amount of moisture inland as they did +previously. In fact, the Andean revolution, even if it occurred as late +as Late Tertiary, would have had no perceptible influence on the +amount of water precipitated on the more eastern parts of the continent. +Oliveira and Leonardos (1943:617) point out that after a Cretaceous +submersion of the central part of Brazil, there was a general uplift. +The authors (_op. cit._:689) mention the presence of continental +Cretaceous deposits in the Central Plateau of Brazil, in support of +these changes, and state that "pelo menos em certas zonas do litoral a +elevação do continente prolongou-se até o Pleistoceno." + +Berry (1942:373) concluded, among other things, that there was a +southward extension "in South America of equatorial floras in the lower +Miocene," and (_op. cit._:372) that ... "east of the Andean Axis in +the south temperate zone there was a normal mesophytic flora ... instead +... of present day large steppes." + +My idea is that a tropical forest still covered the Central Plateau of +Brazil in (early?) Pleistocene times and that populations of +_Proechimys_ of a primitive type, similar to _P. g. steerei_, for +example, lived in that extensive forest-climax. The gradual uplift of +the plateau, however, gradually brought about drier conditions in this +region. As a result a large cliseral change was initiated, which shifted +the forest-climax to the more humid eastern escarpments and lowlands +that were gradually being developed, while the savanna climax was being +established on the plateau. Eventually the effect of the decreasing +moisture was locally accentuated by the erosion of the sandstones +(Oliveira and Leonardos, 1943:690) in northeastern Brazil, thus +depriving it of a natural reservoir of rain water. An arid belt was +developed which now constitutes an efficient geographic barrier to the +distribution of many kinds of animals. + +One marginal species may have shifted eastward with the forest-climax +to effect the Recent distribution. The eastern species became completely +isolated from the main group, accumulated mutations, and evolved into +the subgeneric type _Trinomys_. The generic trend that gave rise to +_Trinomys_ probably remained more stable as far as supraspecific changes +are concerned. The lack of barriers in the distributional area of the +original group favored the dispersal and submergence of mutations and, +therefore, there was but little further supraspecific evolution. The +speciation in both subgenera finally resulted from gradual +differentiation of varying populations since they show combinations of +the generic biotypes and possess few truly qualitative characters. + +The cliseral changes in the Central Plateau, which developed the dry +belt, a barrier, might explain the evolution of a few more supraspecific +groups of mammals, as indicated by the presence of similar forms in the +Amazonian region and in Southeastern Brazil. Among these _Echimys_ and +_Phyllomys_, in the same family with _Proechimys_, show differences that +are parallel to those observed in _Proechimys_. One of these parallel +changes is the increased lamination of the cheekteeth. Although +_Echimys_, from the Amazonian region, has upper molariform teeth with +the four laminae fused, _Phyllomys_ has the four laminae completely +separated. + +None of the genera known from the Upper Oligocene and Miocene of +Argentine deposits seems to be directly ancestral to _Proechimys_. + + + + +SPECIATION + + +The detection of differences of systematic worth between populations of +animals, represented by skins and skulls, is a step preliminary to +deducing the factors responsible for the differences. Ordinarily the +factors which cause heritable differences have to do with geographic +isolation and adaptation to ecological conditions. When differences in +the structure of the animal are known, a person is led to speculate on +the factors which could cause them. For one thing, does the observed +degree of difference tend to isolate animals possessing the "new" +character from the other animals? It would seem to me that the isolation +once started by one of these differences tends to be accentuated with +time and the difference itself thus then becomes a factor responsible +for further differentiation. + +Whether or not transition from one character to another occurs +gradually, in its geographic expression, and thus whether or not +intergradation occurs between two subspecies, can be ascertained by the +analysis of a series of population-samples appropriately distributed +geographically. If two characters of systematic worth are known to blend +in one part of the geographic range of a subgenus, and if the same two +characters are seen in two other populations, far removed geographically +from each other and without any samples of annectent populations to +provide actual evidence of intergradation, then such intergradation is +to be inferred. + +The available collections of _Proechimys_ mostly were made haphazardly +with the result that there are extensive areas from which no specimens +as yet are available. Thus, actual proof of intergradation is often +lacking in areas where it almost certainly occurs. In some extensive +areas, however, many samples, from relatively regular intervals, have +been available and they provide genuine proof of intergradation. These +instances have served as a guide for estimating whether other samples +should be considered to be full species or instead merely subspecies of +the same species. + +Lack of intergradation in any of the characters may be accepted as the +criterion of full species. Where two populations occupying the same +range (sympatric populations) show different qualitative characters, +they almost certainly do not crossbreed. Furthermore the characters that +distinguish such kinds of nonintergrading animals are likely to be +considered as of full specific value when detected in far distant parts +of the range of the subgenus. + +In a genus that is widespread and continuously distributed, it is useful +to know which characters always distinguish full species and which ones, +sometimes or always, distinguish only subspecies, since in a population +from a small island, there is, ordinarily, less individual variation +than in a corresponding population from the mainland or a larger island; +under certain circumstances a person might be tempted to give specific +rank to the population when its characters actually are analogous to +those separating subspecies elsewhere. + +Sometimes it is convenient to recognize species-groups, a systematic +category without nomenclatural status, intermediate between the species +and the subgenus. When there are two groups of species not sharply +separated, including one species whose characters overlap those of each +of the two groups, it would seem most appropriate to recognize only +species-groups instead of subgenera. When, on the other hand, the two +groups of species have mutually exclusive characters and a species with +intermediate characters is unknown, the two groups of species can +conveniently be accorded separate subgeneric rank. + + +SUBGENERIC VARIATION + +A few characters are common to one group of species and other features +are common to a second group. The most striking of these features is the +character of the main fold in the molariform teeth. In one group the +fold transversely crosses the crown of the tooth and in the other it +extends scarcely halfway across. No specimen is intermediate in this +respect. These two groups, furthermore, are separated geographically by +an important barrier, the arid belt that starts in the northeastern +littoral of Brazil (Ceará), and that extends south and southwesterly, +more or less accompanying the São Francisco River in the Plateau, to +about 20° S. _Proechimys_ is thought not to inhabit this arid belt. At +the latitude of 20° S the conditions become more suitable for +_Proechimys_, especially along the rivers which flow eastward, but there +the Plateau is replaced by mountains: the Serra Geral at the west, and +Serra da Mantiqueira at the south; these ranges are bare of forests at +higher elevations. Two groups of species of _Proechimys_ are, therefore, +kept geographically isolated: one group lives in southeastern Brazil, +and the other lives in a large area to the west which starts at 21° S in +Paraguay and Brazil and widens northward and includes, farther west, +central and northern Brazil and all the South American countries +above 21° S, as well as Central America northward to southern Nicaragua. + +The two groups which are here treated as subgenera may be designated as +follows: + +_Trinomys_--main fold deep: aristiforms well-developed on the rump and +outer thighs; tail no less than 75 per cent of length of head and body; +skull without ridges across the parietals; no conspicuous groove for +transmission of nerve inside infraorbital foramen; molariform teeth +decreasing in size from premolar to third molar; 1 to 3 counterfolds in +the molariform teeth. + +_Proechimys_--main fold shallow: aristiforms not developed on rump and +outer thighs; tail less than 75 per cent of length of head and body; +groove for transmission of nerve present in infraorbital foramen of +several subspecies; molariform teeth increasing in size from premolar to +second molar; 2 to 5 counterfolds in molariform teeth. + +Most of these characters vary but do not overlap. Subgeneric rank is +here accorded to the two groups of _Proechimys_ characterized +immediately above. + +The primary cause of the subgeneric differentiation is thought to have +been geologic changes in the continental area. As already pointed out +(see Paleontology), decreasing humidity in the Central Plateau of Brazil +may have caused a migration southwestward of one or more of the species +along with the forests. Once isolated geographically, the species +probably differentiated at an accelerated rate. + +The fact that a much larger number of subspecies occupies the larger +geographic range of the subgenus _Proechimys_ would not be sufficient to +prove that this subgenus, _Proechimys_, is nearer to the primitive group +than _Trinomys_, the subgenus occupying the smaller range with fewer +subspecies. The paleontological evolution of the rodents, however, +consistently points to teeth with a larger number of counterfolds (as +seen in _Proechimys_) as the primitive condition. The extension of the +main fold, tending to set apart one lamina in each upper molariform +tooth, seems to be a specialization; reduction in the size of the head +and body, increase in length of tail and decreasing size of molars +posteriorly also may be specializations. The main point, however, is to +establish if _Trinomys_ is a relic group rather than a "differentiated" +one. If an intermediate form were known which connected _Trinomys_ with +one species of _Proechimys_ more than with another or even if _Trinomys_ +itself more closely resembled one of the groups of species of the +subgenus _Proechimys_ than it did another, we would assume that +divergence and selection accounted for the subgeneric variation. The +lack of any such connecting link favors the first idea, namely that +_Trinomys_ differentiated rapidly with the aid of geographic variation. + +If _Trinomys_ is, as I am inclined to consider it, the result of +"differentiation," its subgeneric features are to be admitted as "new" +and therefore the most primitive species in the genus should be found in +the subgenus _Proechimys_. + +It is a matter of common sense to admit the two groups considered above +as subgenera rather than genera. Since the two structural plans were +established they would, and do, act as different sources of variation. +On the other hand, the morphological differences do not give the two +groups an amount of morphological differences that would justify full +generic rank for each. + + +SPECIFIC VARIATION IN THE SUBGENUS PROECHIMYS + +Most of the described forms in the subgenus were initially named as +distinct full species. More recently, however, in accordance with the +ideas now prevalent in systematic work, many of the named kinds were +reduced to the rank of subspecies. Tate first made a geographic +arrangement (1935:399-400) and later (1939:177-178) provisionally +synonymised several named kinds of _Proechimys_ with _Proechimys_ +"_cayennensis cayennensis_." A similar tendency was clearly displayed by +Ellerman (1940:115-122) who allocated 29 names, out of 33 (in the +subgenus, as here understood), to the species _Proechimys guyannensis_ +and gave full specific rank to four other named kinds. Osgood (1944) +also had the same viewpoint; that is to say, he appeared to have the +idea that there were only two full species in the subgenus in +Brazil--admitting this orally--and consequently he synonymised some full +species where two or more occurred in the same place, thinking that he +was dealing with individual, rather than specific, differences. +Evidently the number of species in the subgenus cannot be great because +the known kinds show few patterns worthy of specific designation and +therefore the majority of the existing names should be suspected of +having no more than subspecific value. Nevertheless none of the above +writers presented real evidence in support of his arrangement. + +Criteria for the recognition of full species are most easily recognized +where two or more different species live together. In the literature, +_P. goeldii_ and _P. "oris"_ were mentioned by Thomas (1912:89) as +having been collected in the same place; _P. mincae_ and _P. +canicollis_, by H. H. Smith (in Allen, 1904:440); _P._ "_leucomystax_," +from Utiarití, by Miranda Ribeiro (1914:42) and _P._ "_longicaudatus_," +from the same place, by Allen (1916:569) were other examples. In these, +and other alleged instances of two or more kinds occurring together, +detailed study of the specimens concerned was necessary to learn the +true facts. Also with the opportunity to compare collections from +several different places, new facts emerged. _P. longicaudatus_, as it +was conceived of by Allen, was a composite species, but in one locality, +Utiarití, Ribeiro and Allen actually were dealing with two distinct +species. + +The species, or subspecies belonging to different species, living +together are: _goeldii_ and _hyleae_, at Fazenda Paraiso; _goeldii_ and +_riparum_ in Manaus; _boimensis_ and _hyleae_ in Tauarí; _leucomystax_ +and _villicauda_ in Utiarití; _mincae_ and _canicollis_ in Bonda; +_gularis_ and _hendeei_ on the banks of Rio Napo ("same trap lines," +according to P. Hershkovitz, _In Litt._). Study of samples of the above +named pairs of kinds of _Proechimys_ showed the following specific +differences: _goeldii_ is large with narrow aristiforms, has a large and +strongly built skull, with four counterfolds in one or more upper +molars: _hyleae_ is smaller, has wide aristiforms, smaller skull with +less pronounced ridges, and never has more than three counterfolds in +the upper molariform teeth; _riparum_ closely resembles _hyleae_; +_boimensis_ has thin aristiforms, small skull and no more than three +counterfolds in the upper molariform teeth in contrast to _hyleae_, +already discussed; _leucomystax_ closely resembles _boimensis_; +_villicauda_ closely resembles both _hyleae_ and _riparum_; _mincae_ is +similar to _hyleae-riparum-villicauda_; _canicollis_ has the number of +counterfolds in all molars reduced to two; _gularis_ is large, has a +strongly built and ridged skull, some upper molariform teeth with four +counterfolds and wide aristiforms; _hendeei_ closely resembles +_leucomystax_ and _boimensis_. + +The evidence obtained from study of specimens where two or more species +occurred together was applied to the remaining samples and the +geographic distribution was worked out. As a result the arrangement +below was made, including all valid kinds already named and those here +newly named from Brazil. The names of kinds I do not consider as +belonging to the subgenus (and genus) are excluded. These are _Echimys +macrourus_ Jentink, not seen, and _Proechimys cayennensis hoplomyoides_ +Tate (= genus _Hoplomys_). The application of names is tentative, +however, because the types deposited in Europe have not been seen. An +asterisk denotes the forms not seen by me. + + _Proechimys guyannensis: arabupu, arescens, bolivianus, cherriei, + chrysaeolus, guairae, o'connelli, guyannensis*, hyleae, leioprimna, + mincae, nesiotes, ochraceus, oris, poliopus, ribeiroi, riparum, + trinitatis, urichi, vacillator*, villicauda, warreni._ + + _Proechimys longicaudatus: boimensis, brevicauda, elassopus, + hendeei, leucomystax, longicaudatus, nigrofulvus, pachita, + rattinus*, roberti, securus, simonsi._ + + _Proechimys semispinosus: amphichoricus, burrus, calidior, + centralis, chiriquinus, colombianus, decumanus, goldmani*, + gorgonae, gularis, hilda*, ignotus, kermiti, liminalis, + panamensis, rosa*, rubellus, semispinosus._ + + _Proechimys goeldii: goeldii, steerei._ + + _Proechimys canicollis._ + +_Proechimys guyannensis_ appears to be more plastic than any other +species. In size of animal, width of aristiforms, color and number of +counterfolds in the cheekteeth, it shows marked response to variations +in geographic conditions. _Proechimys longicaudatus_ is apparently less +plastic; only the number of counterfolds shows marked variation. +_Proechimys semispinosus_ varies much within its range. _Proechimys +goeldii_ seems to be relatively uniform. _Proechimys canicollis_ shows +relatively little variation throughout its range but probably is +divisible into two or more subspecies. + +The primitive _Proechimys_ probably was large with a short tail, narrow +aristiforms, strongly built skull, and five counterfolds in each +molariform tooth. Primitiveness here is inferred from characters which +now are of general occurrence in the whole group as opposed to those +restricted in geographic occurrence. + +It is a curious fact that in this genus, populations from small islands +are more primitive than populations on the mainland. Apparently a small +population restricted to a small island tends to revert to the primitive +type. The homozygous condition will tend toward a generalized genotype +and the disappearance of secondary biotypes. _P. i. iheringi_ on the +Island of São Sebastião averages larger, has thinner aristiforms, and a +stronger skull than the same subspecies on the mainland, and the +cheekteeth usually have two and three counterfolds. The same subspecies +on the mainland has no more than two counterfolds. _Proechimys +semispinosus gorgonae_ and _Proechimys semispinosus ignotus_, living on +Gorgona and San José islands, respectively, are both characterized by +large size, short tails, strong and conspicuously ridged skulls, and +cheekteeth frequently with four and five counterfolds. On the mainland, +closely related subspecies, like _P. s. panamensis_, _chiriquinus_ and +_gularis_, far less frequently have four counterfolds in more than one +or two teeth. More striking still is the population-sample of _gularis_ +from the island of Llunchi, in the Rio Napo, eastern Ecuador. In it +there is a higher ratio of cheekteeth with four counterfolds than there +is in the samples from the banks of the river. + +The two insular forms, _P. s. gorgonae_ and _P. s. ignotus_, referred to +as primitive in the discussion above, have wide aristiforms, which is +contrary to what would be expected in a primitive _Proechimys_. +Supposing, however, as actually seems to be the fact, that narrowness of +the aristiforms depends on an increased number of genes, we deduce that +the population from the mainland, that gave rise to the populations of +the islands, did not have all of the genes necessary to make the +aristiforms narrow. In fact the subspecies known on the mainland, near +the aforementioned islands, have wide aristiforms. + +Another point which favors the idea that narrow aristiforms result from +an increased number of genes is that, generally, the aristiforms are +narrow in any species whose geographic range is extensive and relatively +uniform. + +_Proechimys goeldii_ is the species which has the largest number of +characters that are judged to be primitive, and it may be the oldest +stock. _P. semispinosus_, _P. longicaudatus_ and _P. guyannensis_ may +have been derived from an early splitting of the genus or they may have +branched off the main stem at different times. _P. canicollis_, however, +seems clearly to be an offshoot of _P. guyannensis_; _canicollis_ shows +greater resemblance to _guyannensis_ than to any other species. _P. g. +vacillator_ is another close relative of _P. guyannensis_ with the +number of counterfolds almost as much reduced as in _P. canicollis_. +Conceivably, _vacillator_ is a full species, but the reduction in number +of counterfolds in the teeth more probably expresses only one extreme of +a gradient, as will be discussed below. + + +SUBSPECIFIC VARIATION IN THE SUBGENUS PROECHIMYS + +In spite of the lack of specimens from areas in which _Proechimys_ +certainly occurs, it is evident that the genus has great plasticity and +that the number of subspecies will be greatly increased as additional +material is studied. Only perfunctory examination of samples from +outside the area of Brazil shows me that there are several unnamed +subspecies there. My impression is that Allen's _trinitatis_, of +Trinidad, the genotype of _Proechimys_, will eventually be split. + +There are two main lines of subspeciation in _Proechimys guyannensis_. +The one south of the Amazon River includes _P. g. bolivianus_, in +Bolivia, _P. g. villicauda_, and _P. g. ribeiroi_ occurring on the +divide of the headwaters of the Amazon and Parana rivers, in Brazil, and +_P. g. hyleae_ in the lower Tapajoz and _P. g. nesiotes_ in the lower +Tocantins. All six subspecies have a large number of counterfolds in the +molariform teeth. In these six subspecies, p4 has four counterfolds and +the lower molars have three each. Toward the northeastern coast the +number of counterfolds decreases to three in p4 and to two in the lower +molars, as in _P. g. arescens_, _P. g. leioprimna_ and _P. g. oris_. + +In northern South America, north of the Amazon River, the subspecies +with the greatest number of counterfolds is _P. guyannensis warreni_ +(known from only the Demerara River area); p4 has four counterfolds and +the lower molars have three each. The number decreases in all the +adjacent populations: _P. g. guyannensis_, in the Guianas, _P. g. +trinitatis_, and _P. g. urichi_ (going westward from the Guianas to +Venezuela) have the counterfolds reduced to three in p4, but the lower +molars still have the same number of counterfolds, namely, three, +although there is a tendency for them to coalesce; farther west, on the +coast, the number decreases to three counterfolds in p4 and to only two +in the lower molars as in _P. g. guairae_ and _P. g. mincae_. Subspecies +south of the coast show the same reduction of counterfolds, _P. g. +cherriei_ and _P. g. o'connelli_ being examples; _P. g. ochraceus_ and +_P. g. poliopus_ have the reduction carried to the upper molars, M3 +having usually only two counterfolds; _P. g. chrysaeolus_ in the valley +between the Madalena and the Cauca rivers seems to be somewhat isolated +and shows reversion to three counterfolds in the lower molariform teeth; +directly southward of the range of _P. g. warreni_ the number of +counterfolds decreases to three in all lower cheekteeth (population at +Ayan-Tepuy, southern Venezuela), and then to three in p4 and to two in +the lower molars, as in _P. g. arabupu_ on the Brazilian side of Mount +Roraima, and the reduction is extended to the upper molars in _P. g. +vacillator_. + +On the north bank of the Amazon, the only population of _P. g. hyleae_ +known to me (from Obidos) has four counterfolds in p4 and three in the +lower molars; _P. g. riparum_, from Manaus, also on the north bank of +the Amazon, has three counterfolds in p4 and two counterfolds in the +lower molars. _P. g. hyleae_ occurs also on the south bank of the +Amazon. _P. g. riparum_, therefore, may be the northern part of the +southern cline, instead of the southern end of the northern cline. + +The whole picture, as outlined above, may be explained by assuming that +the species _P. guyannensis_ differentiated somewhere on the Central +Plateau of South America, with three counterfolds in each upper +molariform tooth, four counterfolds in the lower premolar and three +counterfolds in the lower molars. The species might have extended its +range to the Guianas and then all the biotypes with reduced number of +counterfolds might have slowly developed by natural selection. The +gradient is, broadly, from subspecies with greater number of +counterfolds in more humid areas, to a gradually lessening number of +counterfolds in less humid areas. + +_Proechimys longicaudatus_ is limited in the south to the headwaters of +the Parana River drainage, where the subspecies _P. l. roberti_ and _P. +l. longicaudatus_ are found. The species ranges northward through the +Tapajoz drainage, with _P. l. leucomystax_ in the headwaters and _P. l. +boimensis_ in the lower course. To the northwest and west the species is +represented in Bolivia by _P. l. securus_; _P. l. elassopus_, _P. l. +simonsi_, _P. l. pachita_, and _P. l. hendeei_ occur in Peru and _P. l. +brevicauda_ in Peru and Brazil; and _P. l. nigrofulvus_ occurs in +Ecuador. Again in _P. longicaudatus_ it seems that the number of +counterfolds follows a gradient from more humid areas with four +counterfolds in p4, as seen in _nigrofulvus_, _pachita_, _simonsi_, +_elassopus_ and _brevicauda_, decreasing to three or four in _securus_, +to three in _longicaudatus_, but with m3 having only two counterfolds in +_leucomystax_ and _roberti_. _P. l. boimensis_, widely separated in the +lower Tapajoz (no samples being known from the intervening range) may be +the end of a cline started by _leucomystax_ with only 2 counterfolds in +m3 and ending to the northward with four counterfolds in m3. Over the +same area the counterfolds in p4 increase from 3 to 4. + +Of _Proechimys goeldii_ I have had inadequate material but there seems +to be a similar gradient in it which may be traced from _P. g. steerei_ +to _P. g. goeldii_. _P. g. steerei_ has four counterfolds in more upper +molars than occurs in the other subspecies. + +_Proechimys semispinosus_ has its wide range in the mountainous, western +area of South America, the headwaters of the Amazon drainage and +northward in Central America and the nearby Pacific Islands. In these +populations a gradient may exist in the number of counterfolds which is +varied in every population. The highest number seems to occur in the +populations from northern Peru and Ecuador, decreasing from there in all +directions, except in the Panamanian and Columbian islands. In gross +examination, it seems that the size of the animals increases to the +northwards. + + +SPECIFIC VARIATION IN THE SUBGENUS TRINOMYS + +Some specific characters are duplicated in each of the two subgenera; +that is to say, there are some parallel developments and they give the +common generic stock its biotypical variability. Among these parallel +developments are the width of the aristiforms, the amount of pigment in +the agouti-colored setiforms, and the shape of the nasal bones. Other +characters, however, appear in one subgeneric group and not in the +other. The specific variation will be discussed separately for each +subgenus. + +The aristiforms are narrow and soft in _P. dimidiatus_ and in the other +species are wide and stiff, and on the outer thighs and rump some are +light-colored. _P. albispinus_ has the maximum number of light-colored +aristiforms; they are present over the sides and back. This species has, +however, a type of aristiforms unique in the genus--the clavate type. +The tail is longer in _P. iheringi_ and _P. setosus_ than in _P. +dimidiatus_ and _P. albispinus_; the longer type is associated with a +penicillate tip suggesting an adaptation to arboreal habit. The skull +and nasals are longer in _P. dimidiatus_ and _P. iheringi_ than in _P. +setosus_ and _P. albispinus_. In the latter two species the longitudinal +dorsal outline of the skull is conspicuously convex as opposed to +slightly convex in the other two species. The palate is longest in _P. +dimidiatus_ and _P. iheringi_ extending posteriorly to the level of the +second molars; it is slightly shorter in _P. setosus_ and shortest in +_P. albispinus_ where it does not extend behind the level of the first +molars. The incisors are opisthodont in _P. dimidiatus_ and _P. +iheringi_ and orthodont in _P. setosus_ and _P. albispinus_ and even +proodont in one part of the last species. + +The molariform teeth have a large number of counterfolds in both _P. +dimidiatus_ and _P. iheringi_, although the number varies but little in +the first species and much in the second. The variation in _P. iheringi_ +decreases in populations of increasingly more northern geographic +distribution; in both _P. setosus_ and _P. albispinus_ the number of +counterfolds is greatly reduced; there is only one in most specimens of +_P. albispinus_. The incisive foramen is small and nearly round in _P. +dimidiatus_, larger and elongate in _P. iheringi_, very narrow and +fissurelike in both _P. setosus_ and _P. albispinus_. + +The characters of _Trinomys_, as briefly outlined above, seem to be the +result of one original species having split first into four species +which provide a gradient for certain characters. Subsequently one of +these four species, _P. iheringi_, split into six subspecies and +another gradient, parallel to the first, and involving the same +characters, is to be seen. + +The interrelationship among the species is evident, not only because +they have the same subgeneric characters, but because the full species +themselves provide successive steps in a stairway of increasing +specialization from _P. dimidiatus_ to _P. albispinus_. + +Morphologically _P. dimidiatus_ and _P. iheringi_ are sometimes +difficult to distinguish, especially on the basis of cranial features. +Nevertheless close attention to the small, nearly round, incisive +foramen of _P. dimidiatus_ versus the larger, more elongate foramen in +_P. iheringi_ will permit separation of the two. However, the two +species live in the same place and one is led to infer that there may be +greater differences in their physiology than in their morphology. In +fact Dr. H. W. Laemmert, from the Serviço de Estudos e Pesquisas Sobre a +Febre Amarela in Brazil, informs me that while _P. dimidiatus_ was +highly susceptible to the virus of yellow fever (18 out of 24 with virus +in circulation), _P. iheringi_ showed a lower rate of susceptibility (3 +out of 25 with virus in circulation). _P. longicaudatus roberti_, +belonging in the other subgenus, showed no susceptibility at all. + +At Teresópolis, Estado do Rio de Janeiro, the two species were found in +two different forests, only a few kilometers apart, but _dimidiatus_ +lived at a higher elevation, where the humidity was remarkably higher. +Naturally the plant associations were different in the two forests. This +seeming ecological adaptation of the two kinds of _Proechimys_ may +explain why _P. iheringi_ ranges farther north; the forests to the +northward are less humid. + +One of the four species, _P. setosus_, subspecies _elegans_, was used by +Winge (1941:80, 82) as representative of the genus _Proechimys_ when he +was estimating the relationships of that genus. Because _Cercomys_, with +four crests in each of its cheekteeth, was, on other grounds, regarded +by him (_op. cit._: 80) as "... the most primitive genus within the +group.", and because he noted in _P. s. elegans_ 4 crests in P4 and in +some first molars, he concluded that _Proechimys_ was "very closely +related to _Cercomys_." His conclusion seems to be correct, but actually +other species of _Proechimys_ (subgenus _Trinomys_), for example, _P. +dimidiatus_, have four or more crests in each cheektooth, and, +therefore, may be considered as more closely related to _Cercomys_ than +is _P. setosus_. If a large number of crests indicates primitiveness, +_P. dimidiatus_, always with four, is more primitive than any other +species in the subgenus _Trinomys_. Also, the large skull, long hind +foot, short tail and thin aristiforms of _P. dimidiatus_, in my opinion, +are primitive characters. + + +SUBSPECIFIC VARIATION IN THE SUBGENUS TRINOMYS + +One of the species of _Trinomys_, _Proechimys iheringi_, is here +subdivided into six subspecies which show a clinal variation. _P. i. +iheringi_, in the southernmost part of the range of the species (Ilha de +São Sebastião), has three counterfolds in the upper cheekteeth of almost +every young specimen but one of these counterfolds, since it is small, +very shallow, and disappears after little wear, is probably in the +process of disappearance; all lower cheekteeth have two counterfolds or, +rarely, m3 has only one. _P. i. bonafidei_ is the next subspecies +northward, where it was collected at 850 m altitude (Fazenda Bõa Fé). +This subspecies still has two counterfolds in all the upper cheekteeth; +only 3 out of 16 specimens fail to have these counterfolds coalesced in +one or more of the teeth. In the lower cheekteeth the coalescence is +evident in 18 per cent of the specimens. _P. i. gratiosus_, from +Floresta da Caixa Dagua (alt. 750 m), geographically is well removed from +_bonafidei_ (more than two degrees north), and no samples were obtained +from the intervening area. It shows such great reduction in the +counterfolds that the existence of intermediate populations is clearly +suggested. Every upper cheektooth of this subspecies has the two +counterfolds coalesced and in 40 per cent of the specimens M3 has only +one counterfold; in the lower cheekteeth 60 per cent of the specimens +have only one counterfold in m3. _P. i. panema_, occurring approximately +100 kilometers to the northward of _P. i. gratiosus_ (lowland form), has +one counterfold in M3 in only 20 per cent of the specimens but the lower +third molar has only one counterfold in 80 per cent of the specimens. In +_P. i. denigratus_, from about 3 degrees north of the range of _P. i. +panema_, the reduction is proportionately greater: P4 now is the only +upper cheektooth with two counterfolds in every specimen; all molars +tend to have only one; p4 has also two counterfolds but all lower molars +have only one. + +The relative size of the tail also varies in a cline from south to +north. Its length is approximately 87 per cent of the length of the head +and body in _P. iheringi_; 88 per cent in _bonafidei_; 99 per cent in +_gratiosus_; 100 per cent in _panema_; and 103 in _denigratus_. + +One of the subspecies, _P. i. paratus_, however, seems to be completely +out of the dental cline. It was collected in the near proximity of the +type locality of _P. i. gratiosus_, at an elevation of 120 m lower. This +subspecies has two counterfolds in all molariform teeth and only one of +the two specimens known shows these counterfolds coalesced in P4 and M1. +The sample, 2 specimens, is too small to be trustworthy; hence it is +impossible satisfactorily to account for the break in the clinal +variation. Conceivably two full species are involved, but I prefer at +present to defer decision on this problem until such time as more +evidence is accumulated. + +_P. setosus_ is poorly represented, both of the available skins being +faded. Furthermore, no type locality is known for the subspecies _P. s. +setosus_. + +_P. albispinus_ has only two known subspecies: _P. a. albispinus_, +living in a region of higher humidity, is slightly the darker and has +subapical zones of the setiforms on the sides Ochraceous-Tawny; _P. a. +sertonius_, living in a much drier region, has the same subapical zone +Ochraceous-Buff. The number of specimens of _P. a. sertonius_ is so few +that no gradient can be detected, even if one exists. + + + + +TAXONOMIC CHARACTERS + + +Size and Proportions of External Parts + +Absolute size of head and body, tail, hind-foot and ear are useful in +distinguishing subgenera and subspecies and to some extent in +differentiating species. + +The length of head and body is large to medium in _Proechimys_ and +medium to small in _Trinomys_. The tail is long to medium in _Trinomys_ +and short in _Proechimys_. The longest tail, 242 mm, is found in _P. i. +denigratus_, and the shortest tail, 123 mm, in _P. g. steerei_. The +relative length of tail also provides gradients or clines. + +In every species, males surpass females in average size. Nevertheless, +the largest animals are usually females. How this paradoxal fact is to +be accounted for, I am not sure, but it may be that the animals grow as +long as they live and that females have more chances to survive longer +since the care of the young keeps them closer to shelter. + +_Color._--Upper parts vary from Buckthorn Brown to Ochraceous-Buff. Dark +color ordinarily is correlated with an environment of higher degree of +humidity and light color with lower humidity. However, species may be +found in similar conditions of humidity but differing in color. +_Proechimys albispinus albispinus_, for example, a light-colored form, +is found in areas where the rainfall averages 1,000 to 1,500 mm of +annual precipitation, in the isohygra of 80 per cent relative humidity. +These conditions actually are similar to those where _P. dimidiatus_, of +darker color, is found. The subspecies _albispinus_, however, ranges +mostly over a dry area and the fact that it occurs also in a moist area +without appreciable change in color is difficult to explain. + +Insular populations are usually darker or richer in color than +corresponding continental populations. On a small island, uniformity of +environment and inbreeding may be responsible for an accumulation of +characters for richness of color. + + +Pelage + +The pelage provides most useful taxonomic characters. Excepting the +vibrissiform hairs, all of the elements of the pelage have a common +feature, the flattened shape. The hair constellation (_cf._ Toldt, 1935) +on the upper and lateral surfaces is composed of hairs of two main +types: _aristiforms_ (guard hairs) and _setiforms_ (over hairs). + +The aristiforms are wide, strong, and have the dorsal (= anterior) +margins raised, forming a wide shallow longitudinal groove on the dorsal +face of the hair. The tip is a filament that usually is lacking in +aristiforms which are especially strong. Wear probably removes these +tips. The aristiforms have the bases whitish or grayish and the amount +of pigment gradually increases distally to a dark brown or blackish +shade. On the dorsal and lateral surfaces of the head the aristiforms +are small and narrow but gradually increase in length and width caudad +on the animal. The maximum development is reached in the middorsal +region, from where they decrease in size and number toward the lateral +surfaces or caudad. This decrease in the development of the aristiforms, +however, is not uniformly gradual. Generally, the aristiforms become +increasingly conspicuous in a middorsal band, but they extend to the +sides and onto the outer sides of the thighs; the band narrows rapidly +on the rump. In the subgenus _Trinomys_, where the aristiforms attain +their maximum development, they are still strong and conspicuous on the +rump and sometimes around the base of the tail. In _Proechimys_ the +aristiforms do not extend caudad from the hips. Also, in _Trinomys_, +besides the ordinary lanceolate type, there are some aristiforms on the +dorsal surface with a clavate shape; the base is wide and the distal +part narrow. This parallels the conditions in the pelage of the most +spiny species in the genus _Echimys_, _Echimys chrysurus_ +(Lichtenstein). + +The recently named subspecies _Proechimys cayennensis hoplomyoides_ +Tate, 1939, shows an extraordinary development of the aristiforms on +the back and sides such as occurs in the genus _Hoplomys_. Actually the +small bulla, wide basisphenoid and tooth structure add to the +possibility of _hoplomyoides_ being a true _Hoplomys_, and worn teeth +might have been responsible for the difficulty which Tate had in +allocating the form to the proper genus. However, the narrow braincase +is more nearly like that of _Proechimys_ than that of _Hoplomys_. The +intermediate nature of _hoplomyoides_ argues for including the genus +_Hoplomys_ as a subgenus of _Proechimys_. + +Species with narrow aristiforms have a rather soft and flexible pelage, +while those with wide aristiforms have harsh, spiny pelage. The +aristiforms vary in width from 0.45 to 1.3 mm, depending upon the +species or subspecies. + +Animals with narrow aristiforms tend to have a more or less uniform +coloration throughout the dorsal parts. The blackish distal parts of the +aristiforms regularly interline the ground color made by the subapical +zone of the setiforms. If, on the contrary, wide aristiforms occur, the +dorsal surface is conspicuously marked by the wide blackish lines among +spots of color formed by the subapical zones of the setiforms. No clinal +variation was detected in width of aristiforms but geographic variation +in width was noted; for example, the subspecies of _P. iheringi_ differ +in this respect. + +The setiforms are narrow and flattened but are without pronouncedly +raised margins. The setiforms are usually bicolored on the dorsal and +lateral surfaces of the animals, with a subapical zone of some +reddish-brown color, like Ochraceous-Orange or Ochraceous-Buff. They are +whitish or gray on the basal parts and gradually blacken toward the tip, +but have a reddish subapical zone. Common exceptions to this pattern are +setiforms without subapical zones; these appear on the dorsal surface +among setiforms which are normal in possessing distinctive subapical +zones. Also there are setiforms without blackened tips on the lateral +surfaces. Due to their relative abundance and subapical color, these +setiforms are responsible for the dominant color on the upper parts. +Like the aristiforms, they are longer and wider in the middorsal region +of the animal and are gradually less developed on the remainder of the +upper parts. Actually there is more than one type of setiform in the +hair constellation; they vary in length, width and color. Attention was +not given, however, to every type of setiform. + +The ventral surface of the body and the inner sides of the legs are +uniformly covered by short setiforms, thinner and more sparsely +distributed on the inner side of the legs. These setiforms are usually +uniformly white in color or, sometimes, the distal parts are buff or +more richly colored. + +Vibrissiforms are scattered on the dorsal and lateral surfaces of the +body, and in penicillate arrangements on the head. They are longer than +the pelage proper, have a nearly circular cross-section and are blackish +in color. + + +Skull + +The absolute size of the skull is proportionate to bulk of the body. The +supraorbital and parietal ridges are especially developed in the _P. +semispinosus_ group, where they extend across the parietals to the +interparietals. In all members of the subgenus _Proechimys_, these +ridges extend onto the parietal region. In _Trinomys_, however, they do +not extend so far posteriorly as the parietal, but only onto the +squamosal. + +The rostrum varies from slender to stout. Elongate rostra are common in +_Proechimys_; _Trinomys_ has a short blunt rostrum. + +The infraorbital foramen commonly has a ventral groove for nerve +transmission in many forms of _Proechimys_ but _Trinomys_ almost always +lacks this groove. Presence or absence of the groove is a subspecific +character in the subgenus _Proechimys_. + +The jugals are dorso-ventrally wide in _Trinomys_ except in the species +_P. setosus_. In _Proechimys_ a dorso-ventrally narrow jugal is the +rule, but _P. canicollis_ has an especially wide jugal. A postorbital +process appears on the jugo-squamosal suture and is here called +postorbital process of the zygoma. In _Proechimys_ it is more or less +weakly developed and shows no variation of systematic worth. In +_Trinomys_, on the other hand, this process varies in a clinal way (_P. +iheringi_) and stages of the gradient characterize populations of +subspecific rank. + +Linear and spatulate shape of the humular process of the +pterygoid constituted specific characters for Thomas, but there is so +much individual variation in the shape of this process in almost every +population that it has not been used in the present account. + +The mesopterygoid (interpterygoid) fossa in almost every specimen +extends anteriorly to the level of M1 or M2 in _Trinomys_, and to M3 in +_Proechimys_. Exceptions may occur, as in _P. hendeei_, where the fossa +extends to the level of M2. + + +Incisive Foramen + +The shape and dimensions of the incisive foramen long have been +recognized as providing specific characters. Large size of the foramen +is probably correlated with the requirement for a large amount of +moisture reaching Jacobson's organ in the nasopalatine space; the +moistening of the sensory epithelium is certainly involved. There seems +to be a certain correlation between small size of the incisive foramen +and high degree of humidity in the environment. Shapes and dimensions of +the foramen appear as simple or multiple biotypes and provide characters +which can be employed to differentiate subspecies, species and even +subgenera. Usually a character, say a general shape, occurs in nearly +all populations of a given subspecies but the particular shape seems to +be more closely correlated with ecological conditions, especially +humidity. Animals which live far away from large rivers usually have +larger foramina than animals which live close to rivers. + +Both the premaxilla and the maxilla develop processes which form a +sheath for the vomer. This vomerine sheath forms a bridge which +longitudinally crosses the incisive foramen; the structure of this +bridge varies widely. Sometimes the maxillary part is not developed and +the sheath is incomplete posteriorly; sometimes this maxillary part is +very slender and merely touches the premaxillary part. The premaxillary +part, however, is always well developed. + + +Teeth + +Considered by itself the variation in the tooth pattern can lead to +erroneous conclusions as to differentiation of species, because the +number of folds on the occlusal face of a tooth and the depth of certain +folds may be subject to great individual variation as shown by +examination of more than one large series of specimens of the same kind, +age and sex from a single locality. Also there are geographic gradients +or clines, in number of folds. Nevertheless the variation in number of +folds, when measured at sufficient intervals along a cline, may provide +quantitative characters useful in differentiating subspecies. + +[Illustration: FIGS. 2-17. Second left upper molar of the two subgenera +_Proechimys_ and _Trinomys_. All × 8. Anterior border of tooth is at the +top of each figure (Nos. 9 and 17 excepted). Note especially that main +fold is short in _Proechimys_ and long in _Trinomys_. + + FIGS. 2-9. _Proechimys_ (_Proechimys_) _semispinosus liminalis_, + female, MN no. 6243, Rio Quichito. Fig. 2, unworn crown. Figs. 3-8, + cross sections at 0.5 mm. intervals, showing changes in the main fold + and counterfolds at increasing depths as the tooth was ground down. + Fig. 8 is 3 mm. below surface shown in fig. 2. Fig. 9, posterior view + with proximal end of the tooth open showing basal ends of folds. + Later in life the proximal end closes and three roots are formed. + + FIGS. 10-17. _Proechimys_ (_Trinomys_) _iheringi denigratus_, female, + SEPFA no. 17060, Mata do Ribeirão da Fortuna. Figs. 10-16 + corresponding to figs. 2-8. Fig. 17, posterioventral view with + proximal end of the tooth open and part of walls cut away, showing + basal ends of folds. Later in life, as in _Proechimys_, the proximal + end closes and three roots are formed. ] + +The main fold involves both the occlusal face of the tooth and the side +wall. The counterfolds, which are smaller counterparts of the main fold, +in most instances also implicate the wall of the tooth opposite to that +marked by the main fold, but are to be seen mostly on only the occlusal +face of the tooth. Unerupted teeth with the crowns unworn and other +teeth which had barely broken through the gums were ground down to +permit the making of drawings of the surfaces at different levels. This +study revealed that the main fold is deepest in the wall of the tooth. +The development of the main fold varies in two different ways: in all +samples from southeastern and eastern Brazil it is strongly developed, +deeply grooves the tooth through its crown and, in younger individuals, +completely divides the occlusal surface of the tooth. As use wears down +the crown, the main fold soon becomes separated from the opposite wall +and then gradually shortens toward its basal portion. In the other type, +common to animals of all the remaining part of the range of the genus, +the main fold is rather short, never reaching the opposite wall. In this +case, however, one of the counterfolds usually appears almost opposite +the main fold in such a way that in non-erupted or just-erupted teeth +the main fold and one counterfold may be connected by a shallow groove +that may give the impression of extension of the main fold and, +therefore, lead to false interpretations. Closer examination shows that +the counterfold which apparently meets the main fold is really situated +anteriorly or posteriorly to it, in upper or lower teeth respectively. +One subspecies in the subgenus _Trinomys_ differs from the general +characteristics of the subgenus in sometimes showing a small main fold +in P4 whereas it is large in all other cheekteeth. The structural +differences here mentioned in the main fold were never before +recognized. Therefore, the meaning of "quadrilaminate" pattern, "three +outer folds" or "three inner folds" (of authors) is not consistent +insofar as the two groups are concerned. + +Writers have more or less tacitly admitted three as the usual number of +counterfolds present in the upper molariform teeth. Ellerman, for +example (1940:117), states: "Upper cheekteeth normally with three outer +and one inner folds each, these soon becoming isolated as islands. A few +species, which will be discussed below, vary slightly in pattern." +Thomas (1921:140) erected the subgenus _Trinomys_ on the basis of the +upper molariform teeth having only three laminae. Actually a meticulous +study of widely varying samples shows that the number of counterfolds +may vary from _one_ to _five_, the usual number being three or four. One +of the most important facts to record on this subject is that young +specimens with slightly worn molariform teeth are more apt than either +adult or younger specimens to show the maximum number of counterfolds. +Usually nonworn teeth show rounded crests and valleys of different +depth. For example, it is common to see one continuous groove giving the +impression of a main fold crossing the occlusal surface transversely. +The slightest wear of the occlusal surface, however, shows that really +there are two valleys instead of one. The two are the main fold and one +counterfold. In this case, it would be easy to confuse the two types of +teeth, one with the main fold short and the other with the main fold +extending almost all the way across the occlusal surface. + +Some of the counterfolds are especially shallow and tend to disappear in +an early stage of wear, and adult individuals may have these folds +completely worn away. Advanced wear usually develops a cuplike occlusal +surface with only the remains of the main fold and also remains of one +or more counterfolds represented by small enamel islands (Figs. 2 to +17). In the form _Proechimys iheringi iheringi_, for example, every +tooth shows three counterfolds in the upper molariform teeth of +individuals in which the wear is not advanced. This number, however, is +less in all or part of the molariform teeth of older individuals. + +Adjacent counterfolds may appear to be coalesced in many instances. +Coalescence is more likely to be seen in species where a wider variation +in the number of the counterfolds is involved and it appears as a +gradient in the reduction of the number of counterfolds. + +Of great importance, as a general feature of molariform teeth, is the +relative size as related to the geographical distribution, showing, +again, a natural division in the genus. In all forms of southeastern +Brazil the premolars are larger than the first molars, the first molars +are larger than the second molars, and the second molars are larger than +the third molars. The forms from central and northern Bahia, Brazil, +have the molariform teeth more or less the same size. The forms from the +remaining part of the area occupied by the genus have premolars smaller +than the first molars, the first molars smaller than the second molars, +but the second molars larger than third ones. + + + + +HABITS + + +_P. dimidiatus_ was studied in the field and laboratory. _P. dimidiatus_ +in captivity showed regular diurnal activity, coming out of the nest for +food at intervals. Individuals were fed a cereal mixture and nuts of +small size. The animals usually buried the nuts in the sand of the outer +cage. While holding the nut with the mouth and front feet, the animal +patted the sand rapidly, thus burying the nut, and it then pushed more +sand over the place with the front feet. + +Sometimes the emergence from the nest is followed by a long yawning and +stretching ceremony. The animal spreads the fore and hind legs widely +apart, while the back is curved down and the head and tail turn upward. +Then one of the hind legs is stretched backward and, at the same time, +the mouth is opened widely and the tail is moved in an undulatory +fashion. The operation may be repeated using the other legs, or not. + +_P. dimidiatus_ was regularly found in climax forest. The best shelter +and nesting ground was usually under boulders, commonly not farther than +10 meters from water. The entrance to the nest was kept clean. No more +than two adult animals (male and female) were captured in the same +shelter, and only a few times were young captured in the same place with +adults. Nesting places were located also at the bases of trees and near +fallen logs where litter accumulates. + +Records of animals kept in captivity show that the species _dimidiatus_ +survives more than two years. Specimen MN no. 5448 [M] was adult when +captured by the Serviços de Estudos e Pesquisas sobre a Febre Amarela on +December 5, 1938, and died on January 17, 1942. Therefore, it lived for +more than 1,139 days. + +_Proechimys dimidiatus_, in Rio de Janeiro, as well as _P. i. bonafidei_ +and _P. i. iheringi_ which live in the same region, were found breeding +from September to November and from March to May. _Proechimys +longicaudatus roberti_, in Anapolis, Goiaz, was found breeding from July +to November and from January to March. _P. g. hyleae_ and _P. g. +leioprimna_ in the lower Tapajoz and lower Tocantins rivers, Pará, were +found breeding in January. + +It seems that in the Central Plateau and southeastern Brazil, +_Proechimys_ has two litters per year, one in the early spring and a +second in the late summer. The number of young per litter varies from 1 +to 5, although 2 is the usual number. + + + + +CHANGES WITH AGE + + +_Juveniles._--The animals are born with a thick pelage of thin +aristiforms and thin setiforms. The color is uniformly blackish brown. +The nose, hands, feet, ears and tail are pinkish; P4 and M1 are already +erupted and the second molars are included in the bony alveoli. The +incisors are orthodont; the rostrum is short and the braincase is wide. +The posterior part of the skull is greatly curved dorsally. No change is +noticed in the pelage before the second molars erupt and become +functional. + +_Adolescents._--As soon as the second molars become functional, the +pelage starts molting on the back. The thin aristiforms are still in +place but the aristiforms of the adult pelage may be noticed growing +under them in an oval patch which extends from behind the shoulders +caudad to the hips. At that age the first, agouti-colored aristiforms +appear on the mystacial region, immediately behind the vibrissiforms. +The rostrum gradually lengthens and the braincase appears to become less +inflated. + +By the time the third molars erupt, the aristiforms start showing among +the setiforms which are now changing to agouti color in the same area on +the back, while the thin aristiforms of the juvenal stage disappear. The +agouti setiforms are appearing also posteriorly from the mystacial +region to the sides of the head and neck and, at the same time, on the +frontal region. The patch of glossy aristiforms and setiforms on the +back is sharply differentiated from the dull juvenal pelage of the sides +and rump. In a later stage the area of agouti setiforms on the sides of +the neck extends to the outer sides of the arms and finally reaches the +area on the back where the agouti setiforms were already developed. + +_Adults._--When the third molars become functional, the agouti setiforms +are in place except for those on the upper sides of the neck. The +aristiforms have now extended over their normal area of distribution. As +soon as the third molars show wear, the premolars and first molars have +the counterfolds isolated in the occlusal surfaces as enamel islands. +Wearing gradually isolates all counterfolds. + +_Senile_ individuals.--Progression of wear soon eliminates the signs of +the shallowest counterfolds from the occlusal surface and finally the +tooth is reduced to a short crown with a cuplike occlusal surface +completely filled with dentine. Only the main fold usually remains; it +is more or less reduced in size. + +From the records of the Serviços de Estudos e Pesquisas sobre a Febre +Amarela, the following data for males of _Proechimys longicaudatus +roberti_ were obtained: + + ======================+=========+==========+========+=========+======= + | Number | | Length | | Length + AGE | of | Weight | of head| Length | of + | cheek- | in grams | and | of tail | hind + | teeth | | body | | foot + ----------------------+---------+----------+--------+---------+------- + One day old | 2 | 20.5 | 70 | 53 | 24 + 17 days old | 2 | 26.0 | 110 | 60 | 28 + Juvenile | 2 | 85.0 | 150 | 105 | 39 + Adolescent | 3 | 115.0 | 175 | 120 | 45 + Adolescent (older) | 3 | 180.0 | 195 | 135 | 45 + Adult | 4 | 200.0 | 223 | 158 | 48 + Senile individual | 4 | 360.0 | 230 | 170 | 48 + ----------------------+---------+----------+--------+---------+------- + +The weights and measurements (except for one- and 17-day-old animals) +represent averages of specimens of the different ages. + + +Genus =Proechimys= J. A. Allen + +_Genotype._--_Echimys trinitatis_ Allen and Chapman, by original +designation. + + _Proechimys_ Allen and Chapman, 26 December 1899, Bull. Amer. Mus. + Nat. Hist., 12(20):264, orig. description; Tate, 1935, Bull. Amer. + Mus. Nat. Hist., 68(5):398; Ellerman, 1940, The families and genera + of living rodents, Brit. Mus. (Nat. Hist.), 1:115. + + _General characters._--Muriform echimyids of medium size; pelage + with flattened and lanceolate and sometimes clavate aristiforms, + varying greatly in width and distributed over most of the dorsal + surface from shoulders to hips or base of tail; setiforms also + flattened, evenly distributed throughout; entire ventral surface + and inner sides of legs white or, rarely, invaded by some buffy + color; tail shorter than, equal to, or slightly longer than, head + and body, bicolored and with a few bristles between scales, + sometimes heavily penicillated; feet long and narrow; pollex + rudimentary; hallux shorter than fifth toe; ears wide and long; + mammae 2-1=6. + + _Skull._--Generally elongate and strongly built, with supraorbital + ridges well developed, frequently extending across parietals + toward occipital region; zygomatic arches variable in depth, + always with postorbital process; infraorbital foramen with or + without lower groove for transmission of nerve; incisive foramen + usually large; vomerine sheath complete or incomplete; + mesopterygoid fossa extending forward at least to plane of third + molars; bullae large; angular process of mandible turned upward. + +[Illustration: FIGS. 18-21. Occlusal views of the upper left and lower +right molariform teeth of the two subgenera of the genus _Proechimys_. +Anterior end of the tooth row at the top of drawing. All × 6. + + FIGS. 18-19. _Proechimys_ (_Proechimys_) _goeldii steerei_, sex ?, + USNM no. 105537, "Hyutanaham." Upper teeth at left (fig. 18). + + FIGS. 20-21. _Proechimys_ (_Trinomys_) _dimidiatus_, male, MN no. + 6256, Pedra Branca. ] + + _Teeth._--Incisors opisthodont, orthodont or proodont, not grooved; + upper molariform teeth with a main internal fold and one to five + external counterfolds which usually appear as enamel islands in + worn teeth, these counterfolds barely implicating the lateral wall; + lower molariform teeth with folds as in the upper molariform teeth + except that they are reversed and the number of internal + counterfolds is usually fewer in the molars. + + + + +ARTIFICIAL KEY TO THE SUBGENERA AND SPECIES + + + 1. (_a_) Tail less than 90 per cent of head and body; aristiforms + not evident on outer thighs and rump; skull + with ridges across parietals; size of upper cheekteeth + increasing from P4 to M2; main fold small. + subgenus PROECHIMYS, 2 + + (_b_) Tail 90 per cent or more of head and body; aristiforms + evident on outer thighs and rump; skull with + no ridges across parietals; size of upper cheekteeth + decreasing from P4 to M3; main fold large. + subgenus TRINOMYS, 5 + + 2. (_a_) One or more upper molars with four counterfolds 3 + (_b_) Upper molars with no more than three counterfolds, 4 + + 3. (_a_) Aristiforms wide (more than 0.7 mm). _P. semispinosus_, p. 342 + (_b_) Aristiforms narrow (less than 0.7 mm) _P. goeldii_, p. 338 + + 4. (_a_) Aristiforms wide (0.9 mm or more), or narrow (0.6 + to 0.7 mm) but then with only two counterfolds in + each lower molar _P. guyannensis_, p. 355 + (_b_) Aristiforms narrow (0.5 to 0.65 mm) but with one or + more lower molars having three counterfolds. + _P. longicaudatus_, p. 346 + + 5. (_a_) Aristiforms narrow (0.5 mm) and limber; no differentiated + light-colored aristiforms on outer thighs and + rump; incisive foramen short and widest posteriorly + _P. dimidiatus_, p. 371 + (_b_) Aristiforms 0.6 mm or more and stiff; differentiated + light-colored aristiforms on outer thighs and rump; + incisive foramen elongated and constricted posteriorly 6 + + 6. (_a_) Skull large, more than 50 mm in total length, incisors + opisthodont _P. iheringi_, p. 373 + (_b_) Skull small, less than 49 mm in total length, incisors + orthodont or proodont 7 + + 7. (_a_) No clavate aristiforms, tail with white tip, + _P. setosus_, p. 384 + (_b_) Clavate aristiforms among the ordinary ones, tail + without white tip _P. albispinus_, p. 388 + +[Illustration: FIG. 22. Map showing the distribution of the two +subgenera of the genus _Proechimys_.] + +[Illustration: FIG. 23. Map showing the geographic ranges of four +species of the genus _Proechimys_.] + +[Illustration: FIG. 24. Map showing the geographic ranges of four +species of the genus _Proechimys_.] + + +Subgenus =Proechimys= J. A. Allen + + + _General characters._--Pelage with lanceolate aristiforms limited + to an area on the dorsal surface between the shoulders and the + hips; length of tail less than 90 per cent of length of head and + body; skull with conspicuous ridges; extension of supraorbital + ridges always evident on parietals; infraorbital foramen usually + with separate groove for transmission of nerve; palate usually + extended posteriorly as far as third molars; incisors opisthodont; + molariform teeth with a small main fold, never extended + transversely to opposite wall in occlusal surface of tooth; usually + one counterfold anterior to main fold in upper molariform teeth and + posterior to main fold in lower molariform teeth; premolars usually + smaller than first molars, first molars smaller than second molars + but second molars larger than third molars. + + +=Proechimys goeldii= Thomas + + _General characters._--Size large; tail short; aristiforms narrow + and soft, usually concealed in pelage by setiforms; general color + of upper parts some tint of orange, gradually becoming lighter on + sides with no conspicuous, dark longitudinal band on back; feet + dark; ventral surface of body and inner side of legs white but + sometimes with some buff locally; skull broad and strongly built + but not conspicuously ridged; zygomatic expanse great and rostrum + not elongate; incisive foramen narrow; bullae large and inflated; + upper molariform teeth with three to four counterfolds, M3 + ordinarily with four; lower premolars with four, and molars with + three, counterfolds. + + +=Proechimys goeldii steerei= Goldman + + _Proechimys steerei_ Goldman, Proc. Biol. Soc. Washington, 24:238, + 28 November 1911 (original description); Goldman, 1912, Proc. Biol. + Soc. Washington, 25:186; 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; Osgood, 1944, Zool. Ser. + Field Mus. Nat. Hist., 29(13):204. + + _Type locality._--Hyutanaham, Upper Purus, _Lábrea_, Amazonas, + Brazil. _Type:_ United States National Museum, no. 105535, adult + male; collected in 1901 by Prof. J. B. Steere. + + _Range._--Known only from the type locality and Porto Velho. + + _Diagnosis._--Upper parts Mars Orange on back, grading to + Ochraceous-Tawny on sides; zygomatic breadth narrow; nasals short; + incisive foramen narrow and short; vomerine sheath complete and + thick; upper molars usually with four counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Grayish basally, + gradually blackening toward tip; total length, 16 to 19 mm; + maximum width, 0.5 mm. _Setiforms on middorsal region_: Grayish on + basal third, gradually blackening toward tip but interrupted by a + Mars Orange, subapical zone 1.5 mm long; total length, 16 to 19 + mm; maximum width, 0.06 mm. _Setiforms on outer thighs_: Whitish + basally, gradually blackening toward tip but interrupted by Orange + Rufous or Ochraceous-Tawny, subapical zone; total length, 14 to 16 + mm; maximum width, 0.05 mm. + + +[Illustration: FIGS. 25, 27. _Proechimys goeldii steerei_, sex ?, +"Hyutanaham," USNM no. 105537. × 1.] + +[Illustration: FIGS. 26, 28. _Proechimys goeldii goeldii_, female, AMNH +no. 37488. × 1.] + +[Illustration: FIGS. 29, 30. _Proechimys goeldii steerei_, sex ?, USNM +no. 105537, "Hyutanaham." × 1.] + +[Illustration: FIGS. 31, 32. _Proechimys goeldii goeldii_, female, AMNH +no. 37488, Fazenda Paraiso. × 1.] + + _Skull._--Large and strong; rostrum rather pointed posteriorly; + supraorbital ridges not much expanded and extending across + anterior half of parietals; infraorbital foramen without groove + for transmission of nerve, or groove obsolete; zygomatic arches + slender; postorbital process of zygoma involving mostly squamosal; + incisive foramen short and narrow (4.5 × 2.5 mm) with margins + almost parallel or tapering gradually caudad and extending toward + palate as ridges; posterior margin of incisive foramen + approximately 2.5 mm anterior to premolars; vomerine sheath + complete, with both elements well-developed; mesopterygoid fossa + never extending anterior to middle of M3; bullae large, well + inflated and with shallow grooves. + + _Teeth._--Upper molariform teeth: P4 with three counterfolds; + upper molars with four counterfolds each or, less commonly, three. + Lower premolars with four counterfolds; lower molars with three + each. + + _Comparisons._--From _P. g. goeldii_, _steerei_ differs in: Back + and sides with more reddish; narrower interorbitally and across + zygomata; palatilar length less and nasals shorter; maxillary part + of vomerine sheath thicker; usually four instead of three + counterfolds in M3. + +_Remarks._--This subspecies is clearly related to _P. goeldii_. One +skull from Porto Velho, Rio Madeira, _Guaporé_, Brazil (CNHM no. 21558) +may belong to an unnamed subspecies but is provisionally included here. + +In the field notes of Professor Joseph Beal Steere, an entry for no. 72 +reads: "Big white bellied wood rats x two young found in nest of grass +on the ground with the two young--_much darker_ young female." No. 77 in +his field notes corresponds to the type specimen. + + _Specimens examined._--Total number, 4, from Brazil, as follows: + Amazonas. _Lábrea_, Hyutanaham, 3 (USNM); Territ. Guaporé, Porto + Velho, 1 (CNHM). + + +=Proechimys goeldii goeldii= Thomas + + _Proechimys goeldii_ Thomas, June 1905, Ann. Mag. Nat. Hist., 15 + (ser. 7):587, (orig. descr.); Thomas, 1912, Ann. Mag. Nat. Hist., 9 + (ser. 8):89; Thomas, 1920, Ann. Mag. Nat. Hist., 6 (ser. 9):277, + Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):400; Osgood, 1944, + Zool. Ser. Field Mus. Nat. Hist., 29(13):199. + + _Proechimys cayennensis goeldii_, Ellerman, 1940, The families and + genera of living rodents, Brit. Mus. (Nat. Hist.), 1:121. + + _Type locality._--Santarem, _Santarem_, Pará, Brazil. Type: British + Museum (Nat. Hist.), no. 5.1.25.6, adult female; presented by Dr. + E. A. Goeldi. + + _Range._--Margins of the Amazon, between Jamundá and Tapajoz + rivers. + + _Diagnosis._--Upper parts Ochraceous-Tawny; wide across zygomata; + nasals of moderate length; incisive foramen long and narrow; + vomerine sheath complete but maxillary part slender; first and + second upper molars with four counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally and + gradually blackening toward tip which is extended as long, thin + filament; total length, 22 to 24 mm; maximum width, 0.5 mm. + _Setiforms on middorsal region_: Gray basally, gradually + blackening toward tip but interrupted by Ochraceous-Tawny, + subapical zone 3.3 mm long; total length, 19 to 21 mm; maximum + width, 0.06 mm. _Setiforms on outer thighs_: Whitish basally, + gradually blackening toward tip but interrupted by + Ochraceous-Tawny, subapical zone 3 mm long; total length 14 to 16 + mm; maximum width, 0.04 mm. + + _Skull._--Large and strong; nasals pointed posteriorly; + supraorbital ridges moderately developed and extended caudad + across anterior third of parietals; zygomatic arches strong; + postorbital process of zygoma involving mostly squamosal; incisive + foramen elongate and narrow (5 to 6.5 x 2.3 mm) with margins more + or less parallel and raised to form ridges which extend posteriorly + to within 3 mm of plane of premolars; vomerine sheath complete, + with maxillary part thin and extended caudad as medial crest; + mesopterygoid fossa extending forward as far as posterior faces + of second molars or slightly short thereof; bullae large and + inflated. + + _Teeth._--Molariform teeth large, P4-M3 averaging more than 9 mm + in length. Upper molariform teeth: P4 and M3 with three + counterfolds; M1 and M2 with four counterfolds each. In lower + teeth, p4 with four counterfolds and each molar with three + counterfolds. + + _Comparisons._--Differences from _P. g. steerei_ are given in the + account of that subspecies. + +_Remarks._--Specimens from the type locality were not available. +Specimens from Fazenda Paraiso, Faro, were relied upon as representative +of the subspecies. These agree with the type according to Thomas +(1912:89). However, the skin of the type was changed in color by +preservative (Thomas, 1905:587) and the best skin he saw was from Faro +(1912:89). + +Thomas (1920:277) applied the name _goeldii_ also to specimens from +Manacaparú, a place a short distance above Manaus on the Solimões +(Amazon) River and from Acajutuba, near Manaus, on the Negro River. In +referring to these specimens (2 from Manacaparú and 2 from Acajutuba) +Thomas (_loc. cit._) said "Five molar laminae are frequently, if not +invariably, present among these specimens." He did not, however, mention +whether or not the number of laminae was constant in both M2 and M3. One +specimen from Acajutuba, in the collection of Museu Nacional (MN no. +1973 [M]), actually has five laminae in M3, but the specimens in the +American Museum from Faro agree absolutely with Thomas' original +description of _goeldii_. + +Osgood (1944:199) doubted that _goeldii_ was a valid species. Evidence +that Osgood's doubt was unjustified is furnished by the fact that Thomas +(1912:89) pointed out that his specimen from Faro agrees with the type. +Likewise, my two specimens from Faro agree with the type insofar as it +has been described. Thomas (1912:89) mentioned two additional skulls +from the type locality which, he stated, agree with the type which was +received from the Museu Goeldi, Pará. + + _Specimens examined._--Total number, 4, from Brazil as follows: + Pará, _Faro_, Faro, Fazenda Paraiso, 2 (AMNH); Amazonas, _Manaus_, + Manaus, 1 skull (AMNH); Amazonas, _Manaus_, Acajutuba, 1 (MN). + + _Additional records._--Total number, 7 (British Museum), from + Brazil, as follows: Pará, _Santarem_, Santarem (Thomas, 1912:89; + 1920:277), 3; Amazonas, _Manaus_, Acajutuba (Thomas, 1920:277), 2; + _Manacaparú_, Manacaparú (Thomas, 1920:277), 2. + + +=Proechimys semispinosus= (Tomes) + +[Illustration: FIGS. 33, 36. _Proechimys semispinosus liminalis_, +female, MN no. 6253, Rio Quichito. Type. × 1.] + +[Illustration: FIGS. 34, 37. _Proechimys semispinosus amphichoricus_, +male, AMNH no. 77020, Mount Duida. Type. × 1.] + +[Illustration: FIGS. 35, 38. _Proechimys semispinosus kermiti_, female, +AMNH no. 37124, Lower Rio Solimões. Type. × 1.2 (from photograph).] + + _General characters._--Size large; tail short and hairy; + aristiforms wide and stiff, especially well-developed on back; + general color on upper parts some shade of ochraceous, usually much + darker on back and forming a conspicuous dorsal band; feet dark; + ventral surfaces and inner sides of legs white; skull elongate and + strong with ridges well developed; incisive foramen long and + narrow; bullae large; usually four counterfolds in M3 and M2; + usually three but sometimes four counterfolds in M1 and even P4; + lower premolar with four and lower molars with three counterfolds. + +[Illustration: FIGS. 39, 40. _Proechimys semispinosus liminalis_, +female, MN no. 6253, Rio Quichito. Type. × 1.] + +[Illustration: FIGS. 41, 42. _Proechimys semispinosus amphichoricus_, +male, AMNH no. 77020, Mount Duida. Type. × 1.] + +[Illustration: FIGS. 43, 44. _Proechimys semispinosus kermiti_, female, +AMNH no. 37124, Lower Rio Solimões. Type. × 1.2 (from photograph).] + + +=Proechimys semispinosus liminalis= subspecies nova + + _Type locality._--Rio Quichito, affluent from the south of the + Javarí River, near _Benjamin Constant_, Benjamin Constant, + Amazonas, Brazil. _Type_: Museu Nacional, no. 6253, adult female, + collected in August, 1942, by E. Parko. + + _Range._--Known only from the type locality. + + _Diagnosis._--Color uniformly dark, setiforms marked with + Ochraceous-Tawny; skull wide across zygomata; nasals short; + prepalatilar part of skull long; incisive foramen long and narrow; + vomerine sheath incomplete or complete; M2 and M3 almost always + with four counterfolds; M1 more rarely with four counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally, + gradually blackening toward tip which is generally extended as a + filament; total length, 21 to 23 mm; maximum width, 0.9 to 1 mm. + _Setiforms on middorsal region_: Gray basally, gradually + blackening toward tip but interrupted by Ochraceous-Tawny, + subapical zone 3 mm long; total length, 22 to 24 mm; maximum + width, 0.06 mm. _Setiforms on outer thighs_: Whitish basally, + gradually blackening toward tip but interrupted by Ochraceous-Buff, + subapical zone 2.5 mm long; total length, 13 to 15 mm; maximum + width, 0.08 mm; some with gray base, blackening gradually toward + tip, without any subapical zone; some with Light Ochraceous-Tawny, + subapical zone. + + _Skull._--Large and strongly built throughout; supraorbital ridges + expanded and thick, extending, in old specimens, across parietals + to anterior angles of interparietals; interparietal ridges always + conspicuous; rostrum elongated; nasals blunt posteriorly; + zygomatic arches strong; infraorbital foramen with + weakly-developed groove for transmission of nerve; postorbital + process of zygoma involving mostly squamosal; incisive foramen + averaging 6 x 2.7 mm, widest in middle part and posteriorly + constricted, with raised margins which do not extend across + maxillae as ridges; posterior margin of incisive foramen + approximately 1.5 mm anterior to plane of premolars; vomerine + sheath incomplete or, sometimes, complete but always with + maxillary part slender; mesopterygoid fossa not extending forward + past centers of third molars; bullae moderately developed. + + _Teeth._--Upper molariform teeth: P4 always with three + counterfolds; M1 with three counterfolds in 9 of 10 specimens and + four counterfolds in remainder; M2 with four counterfolds in 7 + specimens, three counterfolds in remainder; M3 with four + counterfolds in 6 specimens, three counterfolds in remainder. + Lower premolar always with four, and molars with three, + counterfolds. + + _Comparisons._--From _P. s. semispinosus_, _liminalis_ differs in: + darker color; wider aristiforms; greater percentage of upper + molars with four counterfolds. From _P. s. amphichoricus_, + _liminalis_ differs in: lighter upper parts of almost uniform + color instead of with conspicuous, blackish, middorsal, + longitudinal band; more strongly built skull; longer incisive + foramen; vomerine sheath usually incomplete instead of always + complete. + + _Specimens examined._--Total number, 10 (MN) from the type + locality. + + +=Proechimys semispinosus amphichoricus= subspecies nova + + _Type locality._--Mount Duida, Esmeralda, _Amazonas_, Venezuela; + altitude 325 m. _Type_: American Museum of Natural History, no. + 77020, adult male; collected 16 October 1920 by Olalla Bros. + + _Range._--Headwaters of Negro and Orinoco rivers, along boundary + of Brazil and Venezuela. + + _Diagnosis._--Color dark, blackish on middorsal area; subapical + zone of setiforms on back Buckthorn Brown, but many with distal + parts black; skull broad across zygomata; nasals long; + prepalatilar area of skull long; incisive foramen long and narrow; + vomerine sheath complete; upper molars usually with four + counterfolds but P4 usually with only three. + + _Pelage._--_Aristiforms on middorsal region_: Grayish basally, + gradually blackening toward tip; total length, 18 to 20 mm; + maximum width, 0.8 to 1.0 mm. _Setiforms on middorsal region_: + Gray basally, gradually blackening toward tip but interrupted by a + light (16 i), Buckthorn Brown, subapical zone 2 mm long; total + length, 18 to 22 mm; maximum width, 0.03 mm. Most of them, + however, whitish basally, gradually blackening toward tip without + any distinctively-colored, subapical zone; total length, 24 to 26 + mm; maximum width, 0.5 mm. _Setiforms on outer thighs_: Whitish + basally, gradually blackening toward tip but interrupted by an + Ochraceous-Buff, subapical zone 3.5 mm long; black tip short; total + length, 17 to 19 mm; maximum width, 0.05 mm. + +_Skull._--Large and slender; rostrum elongate; nasals bluntly pointed +posteriorly; supraorbital ridges thick (but not expanded) and extending +across parietals but almost obsolete in middle part of parietals; +infraorbital foramen with weakly-developed groove for transmission of +nerve; postorbital process of zygoma involving mostly squamosal; +incisive foramen 5.5 x 2.8 mm wide in anterior third, with margins +constricted posteriorly and extending as ridges approximately 2 mm +beyond posterior margin of incisive foramen; posterior margin of +incisive foramen approximately 2.5 mm anterior to premolars; vomerine +sheath complete with maxillary part weak and premaxillary part extending +posteriorly beyond middle of incisive foramen; mesopterygoid fossa +extending forward as far as middle of M3; bullae well inflated and +elongated. + +_Teeth._--P4 with four counterfolds in one of five specimens and with +three in remainder; M1 with four counterfolds in three of five specimens +and with three in remainder; M2 with three counterfolds in one specimen +and with four in all four remaining specimens; M3 always with four +counterfolds. Lower premolars with four counterfolds and lower molars +with only three. + +_Comparisons._--The subspecies is easily distinguishable from _P. s. +angularis_ by: larger number of black setiforms on back, forming an +almost black longitudinal band; more elongate skull; larger and longer +bulla; longer incisive foramen which is more constricted posteriorly. + + _Specimens examined._--Total number, 6 (AMNH), as follows: + Venezuela, territ. _Amazonas_, Esmeralda, Mt. Duida, altitude 325 + m., 4; Venezuela, territ. _Amazonas_, Rio Cassiquiare, Quemapuré, + 1; Brazil, Amazonas, _São Gabriel_, Rio Uaupés or Caiari, Tatú, 1. + + +=Proechimys semispinosus kermiti= Allen + + _Proechimys kermiti_ Allen, 30 December 1915, Bull. Amer. Mus. Nat. + Hist., 34(22):629 (orig. descr.); Allen, 1916, Bull. Amer. Mus. + Nat. Hist., 35(30):569; 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._--Lower Rio Solimões (up the Solimões 50 to 60 + miles on the north bank of the river), _Manacaparú_, Amazonas, + Brazil. _Type_: American Museum of Natural History, no. 37124, + adult female; collected 20 April, 1914, by Leo E. Miller (Roosevelt + Brazilian Expedition). + + _Range._--Known only from type locality. + + _Diagnosis._--Upper parts Tawny, with darker longitudinal band on + back, gradually becoming Ochraceous-Buff on sides; zygomata widely + spread; nasals long; incisive foramen long; vomerine sheath + incomplete; only M3 with four counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Grayish basally, + gradually blackening toward tip; total length, 18 to 20 mm; + maximum width, 0.8 mm. _Setiforms on middorsal region_: Grayish + basally, gradually blackening toward tip but interrupted by Tawny, + subapical zone 2 mm long; total length, 18 to 20 mm; maximum + width, 0.06 mm; some blackened toward tip without subapical zone. + _Setiforms on outer thighs_: Whitish basally, gradually blackening + toward tip but interrupted by Ochraceous-Buff, subapical zone + 2.5 mm long; total length, 18 to 20 mm; maximum width, 0.05 mm. + + _Skull._--Large, elongate, and strongly built; rostrum not + conspicuously elongated; nasals bluntly pointed posteriorly; + supraorbital ridges wide and extending posteriorly across parietals + almost to level of interparietal; infraorbital foramen with + moderate development of groove for transmission of nerve; zygomatic + arches slender; postorbital process of zygoma involving mostly + squamosal; incisive foramen 6.5 mm long and 2.7 mm wide, wider in + anterior third and gradually constricted posteriorly, with margins + extended toward palate as ridges; vomerine sheath incomplete, + maxillary part threadlike; mesopterygoid fossa extending forward as + far as anterior third of m3; bullae large and well inflated. + + _Teeth._--P4 with three counterfolds; M3 with four counterfolds; + M1 and M2 with three counterfolds. Lower premolars with four + counterfolds; lower molars with three counterfolds. + + _Comparisons._--From _P. s. amphichoricus_, _kermiti_ differs in: + upper parts Tawny instead of Buckthorn Brown; incisive foramen + longer and wider; vomerine sheath incomplete; only M3 instead of + usually all molars, with four counterfolds. From _P. s. + liminalis_, _kermiti_ differs in: upper parts Tawny instead of + Ochraceous-Tawny; aristiforms narrower; M3 only, instead of + usually M2 and M3, with four counterfolds. + + _Specimens examined._--Only the type. + + +=Proechimys longicaudatus= (Rengger) + + _General characters._--Size medium to large; tail short; + aristiforms long and narrow; general color on upper parts + Ochraceous-Buff to Ochraceous-Orange, finely and uniformly lined + with blackish and not forming evident dark band on back; feet + dorsally white or gray; underparts of body and inner sides of legs + white; skull elongate and slender with moderate ridges; incisive + foramen of medium size; vomerine sheath complete or incomplete; + bullae large and elongate; upper molariform teeth with three + counterfolds; lower molariform teeth with three counterfolds but + commonly one or two molars have only two although premolar may have + four. + +_Remarks._--The identity of "_Echimys longicaudatus_ Rengger" can be +ascertained only after samples have been collected in the area indicated +by Rengger: "unter dem ein und zwansigsten Breitengrade" in Paraguay. Of +the samples available to me, those from Urucum, in western Brazil, are +geographically nearest the type locality. North of Urucum, both in +Brazil and Bolivia, two species of _Proechimys_ live together and one of +them is the same species as that at Urucum. Of the two species found to +the northward in Brazil and Bolivia, the one that ranges farther south +probably will occur at the locality indicated by Rengger. Provisionally, +therefore, the name _longicaudatus_ is allocated to the Urucum sample +(see Osgood, 1944:198). In fact, the lack of a type specimen and the +general nature of Rengger's description make "_Echimys longicaudatus_" a +_nomen vanum_. If two species are found living together in the region of +northern Paraguay indicated by Rengger it probably will be impossible to +be sure to which one his vague description applies. + +The form from Urucum, to which the name _Proechimys longicaudatus_ is +here applied, is undoubtedly closely related to _Proechimys leucomystax_ +Ribeiro, from Utiarití, on the Rio Papagaio and also to _P. roberti_ and +_P. boimensis_, all from Brazil. _P. longicaudatus_ is used as the name +of the species because it is the oldest of the four names. + +[Illustration: FIGS. 45, 48. _Proechimys longicaudatus boimensis_, male, +MCZ no. 30881, Boim. × 1.] + +[Illustration: FIGS. 46, 49. _Proechimys longicaudatus longicaudatus_, +male, AMNH no. 37085, Urucum. × 1.] + +[Illustration: FIGS. 47, 50. _Proechimys longicaudatus leucomystax_, +male, AMNH no. 37509, Tapirapoã. × 1.] + +[Illustration: FIGS. 51, 52. _Proechimys longicaudatus roberti_, male, +MN no. 6233, Pouso Alto, Goiaz. × 1.] + +[Illustration: FIGS. 53, 54. _Proechimys longicaudatus boimensis_, male, +MCZ no. 30881, Boim. × 1.] + +[Illustration: FIGS. 55, 56. _Proechimys longicaudatus longicaudatus_, +male, AMNH no. 37085, Urucum. × 1.] + +[Illustration: FIGS. 57, 58. _Proechimys longicaudatus leucomystax_, +male, AMNH no. 37509, Tapirapoã. × 1.] + +[Illustration: FIGS. 59, 60. _Proechimys longicaudatus roberti_, male, +MN no. 6233, Pouso Alto, Goiaz. × 1.] + + +=Proechimys longicaudatus brevicauda= (Günther) + + _Echimys brevicauda_ Günther, 1 April 1877, Proc. Zool. Soc. London + for 1876, (49):748, fig. 9. + + _Proechimys brevicauda_ Ihering, 1904, Rev. Mus. Paulista, S. + Paulo, 6:422; Osgood, 1914, Zool. Ser. Field Mus. Nat. Hist., + 10(12):168; Thomas, 1924, Ann. Mag. Nat. Hist., 13 (ser. 9):534; + Thomas, 1927, Ann. Mag. Nat. Hist., 19 (ser. 9):553; Thomas, 1927, + Ann. Mag. Nat. Hist., 20 (ser. 9):604; Thomas, 1928, Ann. Mag. + Nat. Hist., 2 (ser. 10):262; Thomas, 1928, Ann. Mag. Nat. Hist., 2 + (ser. 10):292; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68 + (5):399; Osgood, 1944, Zool. Ser. Field Mus. Nat. Hist., 29 + (13):201. + + _Proechimys cayennensis brevicauda_ Ellerman, 1940, The families + and genera of living rodents, Brit. Mus. (Nat. Hist.), 1:120. + + _Type locality._--Chamicuros, Rio Huallaga, Peru. _Type_: British + Museum (Nat. Hist.), no. 69.3.31.7,[F]; _Lectoparatype_: British + Museum (Nat. Hist.), no. 66.1.29.8, from Upper Amazons (E. + Barttet), selected by Thomas (1900:301). + + _Range._--Region of the headwaters and upper courses of the Juruá + and Ucaiali rivers, eastern Peru and northwestern Brazil. + + _Diagnosis._--Upper parts Tawny and blackish without marked + longitudinal band on back; underparts buffy or white; aristiforms + narrow; skull slender; incisive foramen wide; vomerine sheath + complete; molariform teeth with three counterfolds, except p4 with + four and m3 with only two. + + _Pelage._--_Aristiforms on middorsal region_: Blackish basally, + gradually blackening toward tip which is extended as a long + filament; total length, 18 to 20 mm; maximum width, 0.65 mm. + _Setiforms on middorsal region_: Gray basally, gradually + blackening toward tip but interrupted by Tawny, subapical zone 1.2 + mm long; total length, 19 to 21 mm; maximum width, 0.1 mm. + _Setiforms on outer thighs_: Whitish basally, gradually blackening + toward tip but interrupted by Ochraceous-Tawny, subapical zone 3 + mm long. + + _Skull._--Slender but not elongated; nasals tapering posteriorly; + interparietals wide; supraorbital ridges not much extended and + faintly shown across parietals; jugals dorso-ventrally "wide" (3.5 + mm); postorbital process of zygoma weakly developed; incisive + foramen 5.5 x 3 mm, oval, with posterior borders raised to form + ridges which extend toward premolars; vomerine sheath complete, + with maxillary part laterally compressed and extended toward + palate as ridge; mesopterygoid fossa extending forward as far as + third molars; bullae large and well inflated. + + _Teeth._--Molariform teeth with three counterfolds, except p4 + which has four and m3 which has only two counterfolds. + + _Comparisons._--From _P. l. longicaudatus_, _brevicauda_ differs + in: upper parts Tawny instead of Ochraceous-Buff; lower premolar + with four instead of three counterfolds; m3 only, instead of both + m1 and m3, with two counterfolds. From _P. l. boimensis_, + _brevicauda_ differs in: upper parts Tawny instead of + Ochraceous-Orange; aristiforms wider; m3 with two instead of three + counterfolds. + + _Specimens examined._--Total number, 3 (DZ), from Brazil, Amazonas, + _João Pessoa_, Rio Juruá. + + +=Proechimys longicaudatus boimensis= J. A. Allen + + _Proechimys boimensis_ Allen, 24 July, 1916, Bull. Amer. Mus. Nat. + Hist., 35(27):523; 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. + + _Proechimys cayennensis_ Osgood, 1944, Zool. Ser. Field Mus. Nat. + Hist., 29(13):199. + + _Type locality._--Boim, Rio Tapajoz, Santarem, Pará, Brazil. + _Type_: American Museum of Natural History, no. 37486, adult male; + "October 10, 1911 (ex Museu Goeldi)." + + _Range._--Along lower course of Tapajoz River. + + _Diagnosis._--Upper parts Ochraceous-Orange; incisive foramen + posteriorly constricted; mesopterygoid fossa sharply pointed + anteriorly; p4 with four counterfolds, remaining molariform teeth + with three counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally, + gradually blackening toward tip which is extended as a long + filament; total length, 16 to 18 mm; maximum width, 0.5 mm. + _Setiforms on middorsal region_: Gray basally, gradually + blackening toward tip, but interrupted by short (1.5 mm), + Ochraceous-Orange, subapical zone; total length 18 to 20 mm; + maximum width 0.09 mm. _Setiforms on outer thighs_: White basally, + gradually blackening toward tip but interrupted by + Ochraceous-Orange, or Ochraceous-Buff, subapical zone 3.5 mm long; + total length 15 to 17 mm; maximum width, 0.06 mm. + + _Skull._--Elongate and slender; rostrum slender and long; nasals + evenly pointed posteriorly; supraorbital ridges weak and barely + reaching anteriormost part of parietals; zygomatic arches slender; + infraorbital foramen with groove for nerve-transmission obsolete; + postorbital process of zygoma weak and involving mostly squamosal; + incisive foramen 5 mm long and 2.5 to 3 mm wide, oval, with + margins not much extended caudad as ridges; posterior margin of + incisive foramen approximately 2 mm anterior to premolars; + vomerine sheath complete with maxillary part slender; + mesopterygoid fossa sharply pointed anteriorly and extending + forward as far as anterior part of third molar; bullae of medium + size, smooth and more or less triangular in contour. + + _Teeth._--Crown length of upper molariform teeth 7 to 7.5 mm; all + upper molariform teeth with three counterfolds; lower premolar + with four counterfolds; lower molars with three counterfolds each. + + _Comparisons._--From three of the other four subspecies found in + Brazil, _boimensis_ differs in having four, instead of three, + counterfolds in the lower premolars. Differences from _P. l. + brevicauda_ are indicated in the account of that subspecies. + +_Remarks._--The material available from Boim, Tapajoz, is rather poor +and of no great significance. The Museu Nacional has one specimen from +Boim which agrees with the type in the American Museum of Natural +History. The Museum of Comparative Zoology at Harvard College, however, +has one specimen in its collection (MCZ no. 30888 [M]), adult, also from +Boim, in which the color pattern is different although agreement with +the type specimen is shown in cranial characters and width and length of +hairs. This specimen is much darker than the other specimens (of +orange-tint) assigned to this subspecies. However, in other samples of +this species, similarly dark animals have been noted. It seems that the +orange tint is gained only in old age. Another specimen (MCZ no. 30878 +[F]) agrees in all respects with the type of the subspecies but the +locality recorded on the label is Cametá, on the Tocantins River. +Possibly the subspecies has a range so wide as to include Cametá but I +suspect that the data on the label is incorrect as to locality. The +collector marked [M] on the label although the specimen is a [F]. The +mistake might have occurred through labeling of specimens at a time +later than that at which they were collected. The collector was in both +localities on more than one occasion. + + _Specimens examined._--Total number, 5, from Brazil, Pará, as + follows: _Santarem_, Rio Tapajoz, Boim, 3 (AMNH type, MCZ 1, MN 1); + _Porto de Moz_, Tauarí, 1 (MCZ); _Cametá_ (?) 1 (MCZ). + + +=Proechimys longicaudatus longicaudatus= (Rengger) + + _Echimys longicaudatus_ Rengger, 1830, Naturgeschichte der + Saeugethiere von Paraguay, p. 236. + + _Loncheres myosuros_ Lichtenstein, 1832, Darstellung neuer oder + wenig bekannter Säugethiere, pl. 36 and text. + + _Echimys myosuros Is._ Geoffroy Saint-Hilaire, 1840, Mag. Zool., + Ann. 2 (ser. 2):15, 17; Allen, 1899, Bull. Amer. Mus. Nat. Hist., + 12(20):261. + + _Echimys cayennensis_ Pictet, 1841, Mém. Soc. Phys. Hist. Nat., + Genéve, 9:145; Waterhouse, 1848, Nat. Hist. Mammalia, 2:334. + + _Proechimys longicaudatus_ Thomas, 1901, Ann. Mag. Nat. Hist., 8 + (ser. 7):532; Thomas, 1904, Proc. Zool. Soc. London, p. 240; + Allen, 1916, Bull. Amer. Mus. Nat. Hist., 35(30):569; Tate, 1935, + Bull. Amer. Mus. Nat. Hist., 68(5):400. + + _Proechimys cayennensis longicaudatus_ Ellerman, 1940, The + families and genera of living rodents, Brit. Mus. (Nat. Hist.), + 1:121; Osgood, 1944, Zool. Ser. Field Mus. Nat. Hist, 29(13):198. + + _Type locality._--Northern Paraguay ("unter dem ein und zwansigsten + Breitengrade"). _Type_: Apparently no type specimen was + preserved. + + _Range._--Western Mato Grosso, Brazil, and northern Paraguay. + + _Diagnosis._--Upper parts almost uniformly Ochraceous-Buff; + incisive foramen widest posteriorly; vomerine sheath complete; p4 + and m2 with three counterfolds; m1 and m3 usually with two + counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Dark gray, gradually + blackening toward tip that has long filament; total length 20 to + 22 mm; maximum width 0.6 mm. _Setiforms on middorsal region_: + Whitish basally, gradually blackening toward tip but interrupted + by Ochraceous-Buff, subapical zone; blackish tip extended (3.5 mm) + and thin; total length 19 to 20 mm; maximum width 0.06 mm. + _Setiforms on outer thighs_: Whitish basally, progressively + grayish or blackish toward tip but interrupted by Light + Ochraceous-Buff or Ochraceous-Buff, subapical zone; total length + 18 to 20 mm; maximum width 0.06 mm. + + _Skull._--Slender; supraorbital ridge notably raised; bullae + large, elongate, smooth and inflated; jugals narrow; postorbital + process of zygoma of medium size and constructed entirely of + jugal; incisive foramen wide and large (5.5 x 3 mm), being + wider posteriorly than anteriorly and with posterior margins + raised; vomerine sheath complete, maxillary part slender + and laterally flattened; mesopterygoid fossa extending forward as + far as centers, or anterior margins, of third molars; posterior + palatine foramina on plane with posterior faces, or centers, of + second molars. + + _Teeth._--Upper molariform teeth always with three counterfolds. + Lower molariform teeth: p4 and m2 always with three counterfolds, + sometimes the two anterior ones coalesced in m2; m1 with three + counterfolds in one specimen (33 per cent); m3 with only two + counterfolds in all specimens. + + _Comparisons._--Differences from _P. l. leucomystax_ and _P. l. + roberti_ are given in the accounts of those subspecies. + + _Specimens examined._--Total number, 3 (2 AMNH, 1 CNHM), from + Brazil, Mato Grosso, _Corumbá_, Urucum. + + +=Proechimys longicaudatus leucomystax= Ribeiro + + _Proechimys leucomystax_ Ribeiro, May, 1914, Commissão de linhas + telegraphicas estrategicas de Matto Grosso ao Amazonas, Annexo no. + 5. Hit. Nat., Zool., Mammiferos, p. 42, pl. 24 (orig. descr.); + 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._--Utiarití, Rio Papagaio, Diamantion, Mato Grosso, + Brazil. _Type_: Museu Nacional, no. 2212, adult, skull only, + collected on 5 May 1909, by Prof. A. Miaranda Ribeiro, is here + designated _lectotype_. See remarks. + + _Range._--Serra dos Parecís, Mato Grosso, Brazil. + + _Diagnosis._--Ochraceous-Buff, richly lined with blackish, on + upper parts; some setiforms completely blackened distally; + incisive foramen regularly ovoid; vomerine sheath incomplete; + upper molariform teeth and lower premolar with three counterfolds; + m2 with three counterfolds but m1 and m3 usually with two + counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Grayish basally, + gradually blackening toward tip which is extended as a long + filament; total length 21 to 22 mm; maximum width 0.65 mm. + _Setiforms on middorsal region_: Whitish basally, gradually + blackening toward tip but interrupted by Ochraceous-Buff, + subapical zone; some setiforms grayish basally and gradually + blackening toward distal portion without any colored, subapical + zone; total length 20 to 22 mm, maximum width 0.06 mm. _Setiforms + on outer thigh_: White basally, gradually becoming gray toward tip + but interrupted by Light Ochraceous-Buff, subapical zone; some + setiforms gray basally and blackening toward tip, but interrupted + by Ochraceous-Buff, subapical zone; total length 15 to 18 mm; + maximum width 0.08 mm. + + _Skull._--Slender; supraorbital ridges conspicuous; bullae large, + smooth and inflated, with slight, transverse groove; jugals + narrow; postorbital processes of zygomata small and involving only + squamosal; incisive foramen of medium size but narrow (5 x 2.5 + mm), regularly oval and with margins uplifted posteriorly; + vomerine sheath incomplete but maxillary part projecting forward + and sometimes almost reaching premaxillary part; mesopterygoid + fossa reaching forward as far as centers of third molars; + posterior palatine foramina at plane of posterior faces of second + molars or slightly anterior thereto. + + _Teeth._--Upper molariform teeth with three counterfolds. Lower + molariform teeth: p4 and m2 always with 3 counterfolds; m1 and m3 + with 2 counterfolds. + + _Comparisons._--From _P. l. longicaudatus_, _leucomystax_ differs + in: upper parts richly lined, instead of scarcely lined, with + blackish; incisive foramen narrower, and regularly oval instead of + widest anteriorly. + + _Remarks._--_Proechimys leucomystax_ was described mainly on the + basis of the Utiarití specimen, here designated lectotype of the + species. The specimen from the Juina River is younger, as stated + by Ribeiro in his description. Ribeiro mentions the skin of the + specimen from Utiarití as "em muito mao estado" and I presume it + was discarded as it has not been found in the collection of the + Museu Nacional, Brazil. + + _Specimens examined._--Total number, 6, from Brazil, Mato Grosso: + Cáceres, Salto Sepotube, 2 (MN); Cáceres, Tapirapoã, Rio Sepotuba, + 2 (AMNH); _Diamantino_, Utiarití, Rio Papagaio, 1 skull (MN); + _Diamantino_, Rio Juina, 1 (MN). + + +=Proechimys longicaudatus roberti= Thomas + + _Proechimys roberti_ Thomas, December, 1901, Ann. Mag. Nat. Hist., + 8 (ser. 7):531 (orig. descr.); Thomas, 1904, Ann. Mag. Nat. Hist., + 14 (ser. 7):195; Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. + 9):140; Tate, 1935, Bull. Amer. Mus. Nat. Hist., 68(5):400. + + _Proechimys cayennensis roberti_ Ellerman, 1940, The families and + genera of living rodents, Brit. Mus. (Nat. Hist.), 1:121. + + _Type locality._--Rio Jordão, _Araguarí_, Minas Gerais, Brazil; 960 + meters alt. _Type_: British Museum (Nat. Hist.), no. 1.11.3.62, old + male, collected 8 August, 1902, by A. Robert; original number, 705. + + _Range._--Western Minas Gerais and southern Goiaz. + + _Diagnosis._--Color almost uniformly Ochraceous-Orange on upper + parts; setiforms long and narrow; incisive foramen long and wide; + vomerine sheath usually complete; upper molariform teeth and lower + premolar with three counterfolds; m3 with two, and m1 and m2 with + two or three, counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually blackening toward tip; total length, 21 to 23 mm; + maximum width, 0.6 mm. _Setiforms on middorsal region_: Whitish on + basal half, gradually blackening toward tip but interrupted by an + Ochraceous-Orange, subapical zone 6 to 7 mm long; total length 25 + to 30 mm; maximum width, 0.05 mm. _Setiforms on outer thighs_: + Whitish on basal half, gradually becoming gray and then blackish + toward tip but interrupted by wide, Ochraceous-Buff, subapical + zone, the tip being brownish or blackish; total length, 22 to 23 + mm; maximum width, 0.04 mm. + + _Skull._--Slender; supraorbital ridges bent outward and making + sharp angle at point of frontosquamosal suture, then continuing + backward parallel to each other and extending across parietals; + squamosal taking small part in supraorbital ridges; bullae large + (11 x 8 mm), inflated, with shallow depressions; incisive foramen + not especially long but wide (5 x 3 to 3.5 mm), widest in anterior + third and constricted posteriorly; vomerine sheath usually + complete, with maxillary part reduced to slender threadlike + process or, less commonly, missing; mesopterygoid fossa extending + forward as far as centers of third molars; zygomatic arches strong + with jugals of medium dorsoventral width (approximately 3.5 mm); + postorbital process of zygoma weakly developed and involving only + squamosal; posterior palatine foramina on plane of posterior + margins of first molars or slightly anterior thereto. + +[Illustration: FIG. 61. Map showing the geographic ranges of the +subspecies of three species of the subgenus _Proechimys_ in Brazil.] + + _Teeth._--Upper molariform teeth usually with three counterfolds + (92.5 per cent of 42 specimens); M2 with four counterfolds in 5 per + cent of specimens; M3 with two counterfolds in 2.5 per cent of + specimens. Lower molariform teeth: p4 usually with three + counterfolds (97.5 per cent of 39 specimens); rarely with four (2.5 + per cent); m1 with three counterfolds in 58 per cent and two + counterfolds in 42 per cent of specimens; m2 with three + counterfolds in 61 per cent and two counterfolds in 39 per cent of + specimens; m3 always with only two counterfolds. + + _Comparisons._--From _P. l. boimensis_, _roberti_ differs in: wider + incisive foramen; lower premolar with three, and one or two lower + molars with only two, counterfolds, instead of lower premolar with + four and all lower molars with three counterfolds. From _P. + longicaudatus_, _roberti_ differs in: upper parts Ochraceous-Orange + instead of Ochraceous-Buff; incisive foramen wider in posterior + third than in anterior third. + +_Remarks._--This subspecies seems to be adapted to forests of +post-climactic conditions which is probably typical of most valleys and +margins of the rivers in southern Goiaz and western Minas Gerais. It was +found in Goiaz usually in riparian forests with climactic associations +or in some advanced stage of the sere. The animals also make incursions +into nearby open areas or crops of corn. + + _Specimens examined._--Total number, 52, from Brazil, as follows: + Minas Gerais, _Araguarí_, Rio Jordão (effluent of Parnaiba), 960 + meters alt., 2 (1 CNHM, 1 DZ); Goiaz, _Anapolis_, 1010 meters alt., + 38 (MN); Goiaz, _Pouso Alto_, 768 meters alt., 11 (MN); Goiaz, Tio + São Miguel, 2 (MN). + + +=Proechimys guyannensis= (E. Geoffroy) + + _Mus guyannensis_ E. Geoffroy Saint-Hilaire, 1803, Catalogue des + mammifères du Museum d'Histoire Naturelle, Paris, p. 194. + + _Echimys cayennensis_ Desmarest, 1817, Nouv. Dict. Hist. Nat., + Paris, nouv. ed., 10:59. + + _Proechimys cayennensis_ Allen, 1899, Bull. Amer. Mus. Nat. Hist., + 12(20):261, 264; Tate, 1935, Bull. Amer. Mus. Nat. Hist., + 68(5):399; Ellerman, 1940, The families and genera of living + rodents, Brit. Mus. (Nat. Hist.), 1:120. + + _General characters._--Size medium to large; tail usually short; + aristiforms narrow to wide; general color of setiforms on back + ranging from Tawny to Ochraceous-Buff and becoming gradually + lighter on sides; no conspicuous dark longitudinal band on back; + upper parts of hands and feet white to light brown; underparts + white, including inner sides of legs; skull elongate and not + conspicuously ridged; vomerine sheath complete or incomplete; upper + premolar with three counterfolds and molars with two or three; + lower premolar with three or four counterfolds, and lower molars + with two or three. + + +=Proechimys guyannensis villicauda= subspecies nova + + _Type locality._--Tapirapoã, Rio Sepotuba, _Cáceres_, Mato Grosso, + Brazil. _Type_: Museu Nacional, no. 1932, adult male (color faded); + collected on 2 February, 1909, by Prof. A. Miranda Ribeiro; + original number, 788 A. + + _Range._--Serra dos Parecís, headwaters of Paraguai and Tapajoz + rivers. + + _Diagnosis._--Aristiforms wide and stiff; general color on upper + parts Ochraceous-Orange; incisive foramen long; vomerine sheath + incomplete or complete; lower premolar with four counterfolds, + remaining molariform teeth with three counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually becoming gray toward tip, with distal fourth blackish + and ending as a long filament; total length, 22 to 23 mm; maximum + width, 1 mm. _Setiforms on middorsal region_: Whitish on basal + half, gradually blackening toward tip but interrupted by + Ochraceous-Orange, subapical zone 5 mm long; total length, 26 to + 30 mm; width, 0.04 to 0.12 mm. _Setiforms on outer thighs_: Whitish + on basal half, gradually blackening toward tip but interrupted by + Ochraceous-Buff to Ochraceous-Orange subapical zone; total length + 20 to 23 mm; width, 0.03 to 0.18 mm. + + _Skull._--Strong; supraorbital ridges raised and extending across + anterior fourth of parietals; nasals elongate; bullae rounded, + inflated, with shallow grooves; postorbital process of zygoma + weakly developed and constructed entirely of jugal; incisive + foramen elongate and narrow (5.5 x 2.5 mm), posteriorly constricted + with posterior margins elevated above surface of bones; vomerine + sheath incomplete or complete but, when complete, with maxillary + part filiform and delicate; mesopterygoid fossa extending forward + as far as middle, or even anterior, parts of third molars; + posterior palatine foramina on plane of centers, or even anterior + faces, of second molars. + + _Teeth._--Upper molariform teeth with three counterfolds. Lower + molariform teeth: premolar with four counterfolds, molars with + three counterfolds, sometimes with two folds coalesced in center + of tooth. + + _Comparison._--From _P. g. ribeiroi_, and _P. g. bolivianus_, + _villicauda_ differs in wider aristiforms. From _P. g. ribeiroi_, + _villicauda_ further differs in: larger and wider incisive + foramen; vomerine sheath incomplete or complete instead of always + complete and thick. From _P. g. bolivianus_, _villicauda_ differs + in: lower premolars always, instead of rarely, with four + counterfolds, and m3 always with three counterfolds instead of + usually with only two counterfolds. + + _Specimens examined._--Total number, 4, from Brazil, Mato Grosso, + as follows: _Cáceres_, Tapirapoã, 3 (MN); _Diamantino_, Rio + Papagaio, Utiarití, 1 (AMNH). + +[Illustration: FIG. 62. +_Proechimys guyannensis villicauda_, male, MN no. 1932, Tapirapoã. +Type. × 1.] + +[Illustration: FIG. 63. _Proechimys guyannensis ribeiroi_, male, MN no. +1935, Rio Doze de Outubro. Type. × 1.] + +[Illustration: FIG. 64. _Proechimys guyannensis hyleae_, male, MCZ no. +30887, Tauarí. Type. × 1.] + +[Illustration: FIG. 65. _Proechimys guyannensis nesiotes_, male, CNHM +no. 19496, Ilha de Manapirí. Type. × 1.] + +[Illustration: FIG. 66. _Proechimys guyannensis leioprimna_, female, +CNHM no. 19503, Cametá. Type. × 1.] + +[Illustration: FIG. 67. _Proechimys guyannensis oris_, male, CNHM no. +19495, Providencia. × 1.] + +[Illustration: FIG. 68. _Proechimys guyannensis arescens_, male, CNHM +no. 26440, Fazenda Inhuma. Paratype. × 1.] + +[Illustration: FIG. 69. _Proechimys guyannensis riparum_, female, AMNH +no. 143018, Manaus. Type. × 1.] + +[Illustration: FIG. 70. _Proechimys guyannensis arabupu_, male, AMNH no. +75816, Arabupu. Type. × 1.] + +[Illustration: FIG. 71. _Proechimys guyannensis villicauda_, male, MN +no. 1932, Tapirapoã. Type. × 1.] + +[Illustration: FIG. 72. _Proechimys guyannensis ribeiroi_, male, MN no. +1935, Rio Doze de Outubro. Type. × 1.] + +[Illustration: FIG. 73. _Proechimys guyannensis hyleae_, male, MCZ no. +30887, Tauarí. Type. × 1.] + +[Illustration: FIG. 74. _Proechimys guyannensis nesiotes_, male, CNHM +no. 19496, Ilha de Manapirí. Type. × 1.] + +[Illustration: FIG. 75. _Proechimys guyannensis leioprimna_, female, +CNHM no. 19503, Cametá. Type. × 1.] + +[Illustration: FIG. 76. _Proechimys guyannensis oris_, male, CNHM no. +19495, Providencia. × 1.] + +[Illustration: FIG. 77. _Proechimys guyannensis arescens_, male, CNHM +no. 26440, Fazenda Inhuma. Paratype. × 1.] + +[Illustration: FIG. 78. _Proechimys guyannensis riparum_, female, AMNH +no. 143018, Manaus. Type. × 1.] + +[Illustration: FIG. 79. _Proechimys guyannensis arabupu_, male, AMNH no. +75816, Arabupu. Type. × 1.] + +[Illustration: FIGS. 80, 81. _Proechimys guyannensis villicauda_, male, +MN no. 1932, Tapirapoã. Type. × 1.] + +[Illustration: FIGS. 82, 83. _Proechimys guyannensis ribeiroi_, male, MN +no. 1935, Rio Doze de Outubro. Type. × 1.] + +[Illustration: FIGS. 84, 85. _Proechimys guyannensis hyleae_, male, MCZ +no. 30887, Tauarí. Type. × 1.] + +[Illustration: FIGS. 86, 87. _Proechimys guyannensis nesiotes_, male, +CNHM no. 19496, Ilha de Manapirí. Type. × 1.] + +[Illustration: FIGS. 88, 89. _Proechimys guyannensis leioprimna_, +female, CNHM no. 19503. Type. × 1.] + +[Illustration: FIGS. 90, 91. _Proechimys guyannensis oris_, male, CNHM +no. 19495, Providencia. × 1.] + +[Illustration: FIGS. 92, 93. _Proechimys guyannensis arescens_, male, +CNHM no. 26440, Fazenda Inhuma. Paratype. × 1.] + +[Illustration: FIGS. 94, 95. _Proechimys guyannensis riparum_, female, +AMNH no. 143018, Manaus. Type. × 1.] + +[Illustration: FIGS. 96, 97. _Proechimys guyannensis arabupu_, male, +AMNH no. 75816, Arabupu. Type. × 1.] + + +=Proechimys guyannensis ribeiroi= subspecies nova + + _Type locality._--Rio 12 de Outubro, affluent of the Camararé, + _Mato Grosso_, Mato Grosso, Brazil; about 190 kilometers west of + Utiarití; altitude 414 meters. _Type_: Museu Nacional, no. 1935, + adult male (colors faded); collected on 20 June, 1909, by Prof. A. + Miranda Ribeiro; original number _G._ + + _Range._--Known only from the type locality. + + _Diagnosis._--_Aristiforms wide and stiff_; incisive foramen small + and narrow; vomerine sheath complete and thick; p4 with four + counterfolds; remaining molariform teeth with three counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually blackening toward tip; total length, 19 to 22 mm; + maximum width, 0.8 mm. _Setiforms on middorsal region_: Whitish on + basal half, gradually blackening toward tip but interrupted by + subapical zone probably of some tint of ochraceous; total length, + 22 to 24 mm; maximum width, 0.06 mm. _Setiforms on outer thighs_: + Whitish basally, gradually blackening toward tip but interrupted + by probably light ochraceous, subapical zone; total length 14 to + 16 mm; maximum width, 0.03 mm. + + _Skull._--Slender; supraorbital ridges low; bullae ovate with + shallow grooves; postorbital process of zygoma almost obsolete and + involving mostly jugal; incisive foramen short and narrow (4 × 2 + mm), constricted posteriorly and with posterior margins raised; + vomerine sheath complete and thick; mesopterygoid fossa extending + forward as far as posterior faces of second molars; posterior + palatine foramina on plane with centers of second molars. + + _Teeth._--Upper molariform teeth with three counterfolds. Lower + molariform teeth: p4 with four counterfolds; molars with three + counterfolds which sometimes are fused. + + _Comparisons._--From _P. g. bolivianus_, _ribeiroi_ differs in: + aristiforms wider; incisive foramen shorter and narrower; vomerine + sheath complete and thick, instead of complete or incomplete and + not thick; p4 always with four, instead of usually only three, + counterfolds and lower molars always with three, instead of + sometimes with only two, counterfolds in m3. Differences from _P. + g. villicauda_ are given in the account of that subspecies. + +_Remarks._--The name _ribeiroi_ is given in honor of the late Professor +Alipio Miranda Ribeiro, in recognition of his important work in +Brazilian vertebrate zoology. + + _Specimens examined._--Total number, 2 (MN), from Brazil, Mato + Grosso, _Mato Grosso_, Rio 12 de Outubro; altitude, 414 meters. + + +=Proechimys guyannensis hyleae= subspecies nova + + _Type locality._--Tauarí, Rio Tapajoz, _Porto de Moz_, Pará, + Brazil; approximately 87 kilometers south of Santarem. _Type_: + Museum of Comparative Zoology at Harvard College, no. 30887, adult + male; collected on 19 January, 1934, by A. M. Olalla; original + number 7288. + + _Range._--Region of lower Tapajoz River and banks of Amazon up to + the Jamundá River. + + _Diagnosis._--Aristiforms conspicuously wide and stiff; general + color on upper parts Tawny; incisive foramen long and oval; + vomerine sheath complete but with maxillary part slender or, + sometimes, incomplete; p4 with four counterfolds, rarely three; + remaining molariform teeth with three counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually becoming blackish toward tip; total length, 19 to 21 mm; + maximum width, 1.1 mm. _Setiforms on middorsal region_: a. Gray + basally, gradually blackening toward tip but interrupted by wide (5 + to 6 mm) Tawny, subapical zone; some are whitish basally and + gradually become sooty brown toward tip except for same type of + subapical zone (tip only slightly darker than subapical zone); b. + With the same type described above and some completely blackish, + with the base gray; total length 22 to 25 mm; maximum width, 0.1 + mm. _Setiforms on outer thighs_: Whitish basally, gradually + becoming gray and then blackish toward tip but interrupted by long, + Ochraceous-Tawny, subapical zone; tip, itself, blackish brown; + sometimes this type appears with some lighter ones and sometimes + with completely blackish setiforms; total length 16 to 18 mm; + maximum width 0.3 mm. + + _Skull._--Medium in size and slender; cranium narrow and not + increasing much in breadth posteriorly; rostrum stout, laterally + thick, with masseteric crest well-developed; nasals pointed + posteriorly; supraorbital ridges broad but barely extended across + parietals; zygomatic arches strong; postorbital process of zygoma + involving mostly squamosal; incisive foramen long and narrow (5.5 + to 6 × 2.5 mm), oval and extending posteriorly to point only 2 mm + anterior to premolars; vomerine sheath complete with maxillary + part usually slender; mesopterygoid fossa extending forward as far + as centers of third molars; bulla of medium size, well inflated + and with shallow grooves on surface. + + _Teeth._--Upper molariform teeth with three counterfolds. Lower + premolar with four counterfolds or, sometimes (20 per cent of 15 + specimens), with only three; lower molars with three counterfolds. + + _Comparisons._--From _P. g. oris_ and _P. g. nesiotes_, _hyleae_ + differs in: wider aristiforms; general color on upper parts Tawny, + instead of Ochraceous-Orange; vomerine sheath not always complete, + instead of always complete. From _P. g. oris_, _hyleae_ differs + in: p4 usually with four, instead of only three counterfolds and + all molars with three, instead of only lower molars with three + counterfolds. From _P. g. nesiotes_, _hyleae_ differs in: p4 + usually, instead of always, with four counterfolds; color on back + Tawny instead of Ochraceous-Orange; aristiforms wider. + +_Remarks._--This subspecies shows greater variability than any other in +this species. There are two types of coloration. The most common type of +coloration is dark, with Tawny, subapical zones in the setiforms of the +middorsal region and many completely black setiforms; in the other type +the subapical zone is still Tawny but there are no black setiforms. One +specimen from Obidos, on the north bank of the Amazon, completely agrees +in the characteristics of color and skull with the reddish type and +suggests either that there is an extension of the range of the +subspecies along the lower course of the Tapajoz or that there are two +subspecies, in which event the animals from Tauarí are intergrades +between _hyleae_ and an unnamed, tawny-colored subspecies occurring to +the southward. + +Between 13 and 23 January, 1934, A. M. Olalla collected 10 adult +females, 6 of which contained embryos. Three of the females had 2 +embryos each, two had 3 embryos each and one had only 1 embryo. At this +same time and place only ten per cent of specimens obtained were not +fully adult. + + _Specimens examined._--Total number, 21, from Brazil, Pará, as + follows: _Porto de Moz_, Tauarí, right bank of Tapajoz, + approximately 85 kilometers south of Santarem, 20 (19 MCZ, 1 CNHM); + _Obidos_, Obidos, 1 (MCZ). + + +=Proechimys guyannensis nesiotes= subspecies nova + + _Type locality._--Ilha de Manapirí, Rio Tocantins, Pará, + Brazil. _Type_: Chicago Natural History Museum, no. 19496, adult + male; collected on 9 December, 1910, by Dr. Emilia Snethlage; + original number, 12. + + _Range._--Known only from the type locality. + + _Diagnosis._--Aristiforms wide and stiff; general color on upper + parts Ochraceous-Orange; incisive foramen long, with parallel + borders; vomerine sheath complete and thick; p4 with four + counterfolds, remaining molariform teeth with three counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally, + gradually blackening toward tip; total length, 18 to 19 mm; + maximum width, 0.9 mm. _Setiforms on middorsal region_: Gray + basally, gradually blackening toward tip but interrupted by + Ochraceous-Orange, subapical zone 4 mm long; total length 16 to 19 + mm; maximum width, 0.06 mm. _Setiforms on outer thighs_: Whitish + basally, gradually becoming gray and blackish toward tip but + interrupted by Ochraceous-Buff, subapical zone 3 mm long; total + length, 13 to 15 mm; maximum width, 0.03 mm. + + _Skull._--Of medium size; rostrum short; nasals pointed + posteriorly; postorbital ridges extending caudad across anterior + fifth of parietals; zygomatic arches strong; jugal with process in + posterior part of masseteric fossa; postorbital process of zygoma + involving mostly squamosal; incisive foramen elongate, narrow (5 x + 2.3 mm), and parallel sided; posterior margin of incisive foramen + approximately 3 mm anterior to premolars; margins of foramen + raised to form ridges; vomerine sheath complete, of almost uniform + width and set deeply in foramen; mesopterygoid fossa extending + forward as far as centers of third molars; bullae of medium size + and inflated. + + _Teeth._--Upper molariform teeth with three counterfolds; p4 with + four counterfolds; m1-3 with three counterfolds. + + _Comparison._--_From P. g. oris_, _nesiotes_ differs in: + Aristiforms conspicuously wider; incisive foramen shorter and + narrower, with borders parallel instead of posteriorly + constricted; posterior margin of incisive foramen farther from + premolars; p4 with four, instead of three, counterfolds; lower + molars with three instead of two counterfolds. From _P. g. + leioprimna_, _nesiotes_ differs in: Incisive foramen with parallel + borders instead of oval; p4 with four instead of three + counterfolds; m3 always with three instead of two counterfolds. + +_Remarks._--Dr. E. Snethlage mentions the type as having been collected +at night in the forest. + + _Specimens examined._--Total number, 8 (MCZ, CNHM, MN), from + Brazil, Pará, Tocantins River, Ilha de Manapirí. + + +=Proechimys guyannensis leioprimna= subspecies nova + + _Type locality._--Cametá, left bank of Tocantins River, near its + mouth, Cametá, Pará, Brazil. _Type_: Chicago Natural History + Museum, no. 19503, adult female; collected on 21 January, 1911, by + Dr. Emilia Snethlage; original number, 35. + + _Range._--Known only from type locality but probably extending + westward toward Xingú River. + + _Diagnosis._--Aristiforms wide and stiff; general color on upper + parts Ochraceous-Orange, incisive foramen moderately long; oval; + vomerine sheath complete; all molariform teeth with three + counterfolds, except lower, third molar which has only two. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally, + gradually blackening toward tip which is extended as a short + filament; total length, 19 to 21 mm; maximum width, 0.8 to 1 mm. + _Setiforms on middorsal region_: Whitish basally, gradually + blackening toward tip but interrupted by Ochraceous-Orange, + subapical zone 2 to 3 mm long; total length, 17 to 20 mm; maximum + width 0.1 mm. _Setiforms on outer thighs_: Whitish basally, + becoming gradually gray and then blackish toward tip but + interrupted by Ochraceous-Buff, subapical zone; blackish tip + short; total length 13 to 15 mm; maximum width 0.06 mm. + + _Skull._--Of medium size; rostrum relatively short; nasals with + posterior borders rounded; postorbital ridges extending across + anterior fourth of parietals; zygomatic arches moderately strong; + postorbital process of zygoma involving both jugal and squamosal; + incisive foramen of medium length (4 to 5 mm) and narrow (about + 2.5 mm), oval and extending caudad to a plane approximately 2 mm + anterior to premolars; vomerine sheath complete, with premaxillae + forming approximately anterior 3/4 of sheath; maxillary part of + sheath short but well-developed; mesopterygoid fossa extending + forward as far as centers of third molars; bullae of medium size + but well-inflated. + + _Teeth._--Upper molariform teeth with three counterfolds. Lower + molariform teeth with three counterfolds, except third molar which + has only two. + + _Comparison._--From _P. g. oris_, _leioprimna_ differs in: + conspicuously wider aristiforms; shorter and narrower incisive + foramen; lower molariform teeth with three counterfolds (except m3 + with only two), instead of lower molars with only two + counterfolds. Differences from _P. g. nesiotes_ are given in the + account of that subspecies. + +_Remarks._--The paratype was collected in an "igarapé," depression +usually invaded by the river waters; the paratype, collected on 18 +January, 1911, had two large embryos. + + _Specimens examined._--Total number, 4 (2 CNHM, 2 AMNH), from + Brazil, Pará, Cametá. + + +=Proechimys guyannensis oris= Thomas + + _Proechimys oris_ Thomas, September, 1904, Ann. Mag. Nat. Hist., 14 + (ser. 7): 195; Thomas, 1905, Ann. Mag. Nat. Hist., 15 (ser. 7):587; + Thomas, 1912, Ann. Mag. Nat. Hist., 9 (ser. 8):89; Tate, 1935, + Bull. Amer. Mus. Nat. Hist., 68:400; Osgood, 1944, Zool. Ser. Field + Mus. Nat. Hist., 29:199. + + _Proechimys cayennensis oris_ Ellerman, 1940, The families and + genera of living rodents, Brit. Mus. (Nat. Hist.), 1:121. + + _Type locality._--Igarapé-assú, E. F. B., near Belem, Igarapé-assú, + Pará, Brazil. _Type_: British Museum (Nat. Hist.), no. 4.7.4.78, + old male; collected on 6 March, 1904, by Alphonse Robert; original + number, 1818. + + _Range._--Probably most of the region on south bank of Amazon + River, between Tocantins (west) and Gurupí River (south). + + _Diagnosis._--Aristiforms narrow but somewhat stiff; color on + upper parts Ochraceous-Orange; incisive foramen long and wide, + conspicuously constricted posteriorly; posterior margin of incisive + foramen close to plane of premolars; vomerine sheath complete but + maxillary part threadlike; upper molariform teeth and lower + premolar with three counterfolds; lower molars with only two + counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally, + gradually blackening toward tip, which is extended as a filament; + total length, 16 to 17 mm; maximum width, 0.6 to 0.7 mm. + _Setiforms on middorsal region_: Gray basally, gradually becoming + blackish toward tip but interrupted by Ochraceous-Orange, + subapical zone 2 to 4 mm long; total length, 18 to 20 mm; maximum + width 0.06 mm. _Setiforms on outer thighs_: Whitish basally, + gradually becoming blackish toward tip but interrupted by + Ochraceous-Buff, subapical zone; total length, 15 to 16 mm; + maximum width, 0.04 mm. + + _Skull._--Of medium size; supraorbital ridges well developed and + extending across anterior fifth of parietals; zygomatic arches + strong; jugal with masseteric fossa deep and with well-developed + posterior process; postorbital zygomatic process involving mostly + squamosal; incisive foramen long (6 to 7 mm), widest anteriorly + (2.5 to 3.2 mm); but narrowing posteriorly to less than 1 mm and + extending caudad almost to plane of premolars; vomerine sheath + complete but maxillary part delicate and threadlike; mesopterygoid + fossa extending forward as far as third molars or posterior parts + of second molars; bullae large and inflated. + + _Teeth._--Each upper molariform tooth with three counterfolds. In + lower jaw, premolar with three, and molars with only two, + counterfolds. + + _Comparisons._--Differences from the subspecies with adjoining + ranges are given in the accounts of those subspecies. + +_Remarks._--Thomas (1912:89) extended the known range of the subspecies +to Faro, on the Jamundá River, on the left bank of the Amazon, and to +Boim, on the Tapajoz River, as well as to Benevides, E. F. Braganca, +near Belem. It seems to me that the specimens from Faro should be +referred provisionally to _Proechimys guyannensis hyleae_; the specimens +from Boim are "more brightly rufous" (Thomas, _loc. cit._) and could be +referred to Allen's _P. boimensis_, described in 1914, but _P. +guyannensis hyleae_ probably lives in the same place and only an +examination of the specimens, which I have not seen, would permit of +certainly allocating the specimens to their correct species. The +specimens from Benevides are more certainly _P. g. oris_. + +E. Snethlage collected one specimen in a garden (Providencia, E. F. B.). +However, according to the personnel of the Brazilian Health Service, the +animals are strictly forest dwellers although they do make excursions +into more open places. + + _Specimens examined._--Total number, 3, from Brazil, Pará, as + follows: Providencia, E. F. B., approximately 15 kilometers east + from Belem, 1 (CNHM); Tanaquará, near Belem, 1 (MN); Rio Guamá, + near Belem, 1 (AMNH). + + _Additional record._--Brazil, Pará, Benevides, E. F. B., + approximately 100 kilometers north-east of Belem (Thomas, + 1912:89). + + +=Proechimys guyannensis arescens= Osgood + + _Proechimys cayennensis arescens_ Osgood, 12 July 1944, Zool. Ser. + Field Mus. Nat. Hist., 29(13): 198. + + _Type locality._--Fazenda Inhuma, below Santa Filomena, upper Rio + Parnaiba, _Vitoria do Alto Parnaiba_, Maranhão, Brazil. _Type_: + Chicago Natural History Museum, no. 26441, adult male; collected on + 5 August, 1925, by Heinrich E. Snethlage. + + _Range._--Region including the valleys of the Turí-assú and + Parnaiba rivers, Maranhão, Brazil. + + _Diagnosis._--Aristiforms moderately wide and not conspicuously + stiff; general color of upper parts near (15'a) Ochraceous-Orange; + incisive foramen long and wide; vomerine sheath complete or + incomplete; upper molariform teeth and lower premolar with three + counterfolds; lower molars with only two. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually blackening toward tip; total length, 19 to 21 mm; + maximum width, 0.7 mm. _Setiforms on middorsal region_: Whitish + basally or on basal half, gradually becoming gray and then + blackish toward tip, but interrupted by long (5 to 6 mm) subapical + zone near (15'_a_) Ochraceous-Orange; total length, 15 to 16 mm; + maximum width 0.05 mm. _Setiforms on outer thighs_: Whitish on + basal half, gradually becoming gray and then blackish toward tip + but interrupted by Ochraceous-Buff, subapical zone; tip sometimes + not conspicuously darker than subapical zone; total length 18 to + 25 mm; maximum width, 0.03 mm. + + _Skull._--Medium in size, not elongated; nasals pointed + posteriorly; supraorbital ridges strong and thick, extending + caudad across anterior third of parietals; zygomatic arches + strong; postorbital process of zygoma involving only squamosal; + incisive foramen 5 by 2.7 mm, oval and extending caudad to plane + approximately 2 mm anterior to premolars; posterior margins of + incisive foramen not forming a ridge; vomerine sheath complete and + with maxillary part slender and threadlike, or incomplete, in + which event, maxillary part not extended enough to join + premaxillary process; mesopterygoid fossa extending forward + as far as centers of third molars; bulla large and more or less + triangular in its peripheral outline. + + _Teeth._--Upper molariform teeth with three counterfolds each. + Lower premolar with three counterfolds; lower molars with two + counterfolds. + + _Comparisons._--From _P. g. oris_, _arescens_ differs in: Color of + upper parts lighter and more uniform; incisive foramen oval + instead of conspicuously constricted posteriorly; posterior margin + of incisive foramen farther from premolars. + +_Remarks._--One specimen from Turí-assú (MN) has been identified by O. +Thomas as "_P. oris_" (his own handwriting is on the label) and the +subspecies is really closely related to _oris_. + + _Specimens examined._--Total number, 3, from Brazil, Maranhão, as + follows: _Vitoria do Alto Parnaiba_, Fazenda Inhuma (below Santa + Filomena), 2 (CNHM); Alto da Alegria, Turí-assú, 1 (MN). + + +=Proechimys guyannensis riparum= subspecies nova + + _Type locality._--Manaus, _Manaus_, Amazonas, Brazil. _Type_: + American Museum of Natural History, no. 143018, adult female; + collected 6 March, 1943. + + _Range._--Known only from type locality but probably extending + northward and eastward. + + _Diagnosis._--Aristiforms wide and stiff; upper parts + Ochraceous-Tawny; incisive foramen short, wide, and oval; vomerine + sheath incomplete; upper molariform teeth and lower premolar with + three counterfolds; lower molars with only two counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Gray basally + gradually blackening toward tip; total length, 18 to 20 mm; + maximum width, 0.9 mm. _Setiforms on middorsal region_: Whitish + basally, gradually blackening toward tip but interrupted by 2 mm + long, Ochraceous-Tawny, subapical zone; total length 20 to 22 mm; + maximum width, 0.04 mm. Some are whitish basally and gradually + become black toward tip with no subapical zone. _Setiforms on + outer thighs_: Whitish basally, gradually becoming gray and then + blackish toward tip but interrupted by Ochraceous-Buff, subapical + zone 3 to 4 mm long; tip not conspicuously dark; total length, 17 + to 19 mm; maximum width, 0.03 mm. + + _Skull._--Of medium size and slender; rostrum slender; nasals + rounded posteriorly; supraorbital ridges well developed and barely + extended onto anteriormost part of parietals; zygomatic arches + slender; infraorbital foramen with well-developed groove for nerve + transmission; postorbital process of zygoma almost obsolete and + involving mostly squamosal; incisive foramen short and wide (4.5 × + 3.3 mm), oval and with posterior margins raised to form ridges + which extend toward premolars; posterior margin of incisive + foramen approximately 2.5 mm anterior to premolars; vomerine + sheath incomplete, with only short, premaxillary part; + mesopterygoid fossa extending forward as far as anterior + parts of third molars; bullae large and inflated, with more or + less triangular outline. + + _Teeth._--Crown length of well worn P4-M3, 6.8 mm; upper + molariform teeth with three counterfolds each. Lower premolar + with three counterfolds; lower molars with two counterfolds. + +[Illustration: FIG. 98. Map showing the geographic ranges of the +subspecies of _Proechimys guyannensis_ in Brazil.] + + _Comparisons._--From _P. g. oris_ and _P. g. hyleae_, _riparum_ + differs in: Shorter and wider incisive foramen; vomerine sheath + incomplete, instead of sometimes incomplete. From _P. g. oris_, + _riparum_ differs in: Upper parts Ochraceous-Tawny instead of + Ochraceous-Orange; aristiforms conspicuously wider. From _P. g. + hyleae_, _riparum_ differs in: Aristiforms narrower; upper parts + Ochraceous-Tawny instead of Tawny; lower premolars with three, + instead of four, counterfolds; lower molars with two, instead of + three, counterfolds. + + _Specimens examined._--Type only. + + +=Proechimys guyannensis arabupu= subspecies nova + + _Type locality._--Arabupu, Mount Roraima, _Bõa Vista_, Territ. Rio + Branco; about 1540 meters altitude. _Type_: American Museum of + Natural History, no. 75816, adult male; collected by Dr. G. H. H. + Tate on 30 December, 1927; original number, 4716. + + _Range._--Known only from the type locality. + + _Diagnosis._--Aristiforms conspicuously wide and stiff; color on + upper parts dark, near (15'_j_) Ochraceous-Tawny; incisive foramen + widest in anterior third; vomerine sheath complete, sometimes + incomplete; upper molariform teeth and lower premolar with three + counterfolds; lower molars with two counterfolds. + + _Pelage._--_Aristiforms on middorsal region_: Whitish basally, + gradually blackening toward tip; total length, 19 to 22 mm; + maximum width, 1.1 mm. _Setiforms on middorsal region_: Gray + basally, gradually blackening toward tip but interrupted by dark + (15'j) Ochraceous-Tawny, subapical zone 3 mm long; some completely + blackish on distal parts; total length, 20 to 23 mm; maximum + width, 0.07. _Setiforms on outer thighs_: Whitish basally, + gradually blackening toward tip but interrupted by Ochraceous-Buff + or Light Ochraceous-Buff, subapical zone; total length, 20 to 23 + mm; maximum width, 0.07 mm. + + _Skull._--Size medium; nasals pointed posteriorly; supraorbital + ridges prominent and slightly extended caudad onto anterior half + of parietals; groove for transmission of nerve in infraorbital + foramen weakly developed; zygomatic arches strong; postorbital + process of zygoma involving mostly squamosal; incisive foramen + approximately 5.5 × 2.7 mm, widest in anterior third and + constricted posteriorly, with posterior margin about 1 mm anterior + to plane of premolars; vomerine sheath complete, with premaxillary + part expanded and maxillary part notably slender and sometimes + lacking; mesopterygoid fossa in some specimens extending forward + as far as middle parts of second molars; bullae large and + inflated. + + _Teeth._--Upper molariform teeth with three counterfolds each. + Lower premolar with three counterfolds; molars with only two. + + _Comparisons._--From _P. g. warreni_, _arabupu_ differs in: + Narrower aristiforms; narrower incisive foramen; lower premolar + with three instead of four counterfolds; lower molars with two, + instead of three, counterfolds. From _P. g. oris_, _arabupu_ + differs in: Aristiforms wider; posterior margin of incisive + foramen farther from plane of premolars; upper parts dark (15'_j_) + Ochraceous-Tawny, instead of Ochraceous-Orange. + +_Remarks._--The sample is fairly uniform. + + _Specimens examined._--Total number, 6 (AMNH), from Brazil, + Territorio do Rio Branco, Bõa Vista, Mount Roraima, Arabupu; + approximately 1540 m. altitude. + + +Subgenus =TRINOMYS= Thomas + + _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. + +[Illustration: 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). + +[Illustration: 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). + + +=Proechimys iheringi= Thomas + + _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. + +[Illustration: FIG. 104. _Proechimys iheringi iheringi_, female, MN no. +6453, Ilha de São Sebastião. × 1.] + +[Illustration: FIG. 105. _Proechimys iheringi bonafidei_, male, MN no. +6183, Fazenda Bõa Fé. Type. × 1.] + +[Illustration: FIG. 106. _Proechimys iheringi gratiosus_, male, MN no. +4024, Floresta da Caixa Dagua. Type. × 1.] + +[Illustration: FIG. 107. _Proechimys iheringi panema_, female, MN no. +8288, Campinho. Type. × 1.] + +[Illustration: FIG. 108. _Proechimys iheringi denigratus_, male, MN no. +8500, Mata do Ribeirão da Fortuna. Type. × 1.] + +[Illustration: FIG. 109. _Proechimys iheringi paratus_, female, MN no. +4012, Floresta da Capela de São Braz. Type. × 1.] + +[Illustration: FIG. 110. _Proechimys iheringi iheringi_, female, MN no. +6453, Ilha de São Sebastião. × 1.] + +[Illustration: FIG. 111. _Proechimys iheringi bonafidei_, male, MN no. +6183, Fazenda Bõa Fé. Type. × 1.] + +[Illustration: FIG. 112. _Proechimys iheringi gratiosus_, male, MN no. +4024, Floresta da Caixa Dagua. Type. × 1.] + +[Illustration: FIG. 113. _Proechimys iheringi panema_, female, MN no. +8288, Campinho. Type. × 1.] + +[Illustration: FIG. 114. _Proechimys iheringi denigratus_, male, MN no. +8500, Mata do Ribeirão da Fortuna. Type. × 1.] + +[Illustration: FIG. 115. _Proechimys iheringi paratus_, female, MN no. +4012, Floresta da Capela de São Braz. Type. × 1.] + +[Illustration: FIGS. 116, 117. _Proechimys iheringi iheringi_, female, +MN no. 6453, Ilha de São Sebastião. × 1.] + +[Illustration: FIGS. 118, 119. _Proechimys iheringi bonafidei_, male, MN +no. 6183, Fazenda Bõa Fé. Type. × 1.] + +[Illustration: FIGS. 120, 121. _Proechimys iheringi gratiosus_, male, MN +no. 4024, Floresta da Caixa Dagua. Type. × 1.] + +[Illustration: FIGS. 122, 123. _Proechimys iheringi panema_, female, MN +no. 8288, Campinho. Type. × 1.] + +[Illustration: FIGS. 124, 125. _Proechimys iheringi denigratus_, male, +MN no. 8500, Mata do Ribeirão da Fortuna. Type. × 1.] + +[Illustration: FIGS. 126, 127. _Proechimys iheringi paratus_, female, MN +no. 4012, Floresta da Capela de São Braz. Type. × 1.] + + +=Proechimys iheringi iheringi= Thomas + + _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 bonafidei= subspecies 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 gratiosus= subspecies 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 panema= subspecies 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 denigratus= subspecies 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 paratus= subspecies 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. + +[Illustration: 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. + +[Illustration: 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. + +[Illustration: FIGS. 133, 135. _Proechimys albispinus albispinus_, male, +CNHM no. 20409, Macaco Seco. × 1.] + +[Illustration: FIGS. 134, 136. _Proechimys albispinus sertonius_, male, +MN no. 6454, Bonfim. × 1.] + +[Illustration: FIGS. 137, 138. _Proechimys albispinus albispinus_, male, +CNHM no. 20409, Macaco Seco. × 1.] + +[Illustration: 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 CNHM, 1 MCZ), from + Brazil, Bahia, _Andaraí_, Macaco Seco. + + +=Proechimys albispinus sertonius= Thomas + + _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 CNHM, 1 MCZ, 1 USNM); + _Bonfim_, Bonfim, 6 (5 DZ, 1 MN). + + + + +INCERTA SEDIS + + +=Proechimys myosuros= (Lichtenstein) + + _Loncheres myosuros_ Lichtenstein, 1818, Das zoologische Museum der + Universität zu Berlin, (2):18 (_nomen nudum_); Lichtenstein, 1820, + Abhandl. K. Akad. Wissensch., Berlin (1818-1819):192, pl. 1, fig. 2 + (orig. descr.); Wied, 1826, Beiträge zur Naturgeschichte von + Brasilien, 2:445. + + _Mus leptosoma_ Brants, 1827, Het geslacht der Muizen door + Linnaeus opgesteld ..., Berlyn, p. 150; Lichtenstein, 1830, + Darstellung neuer oder wenig bekannter Säugethiere, Berlin, Heft + 7, pl. 36, fig. and text pages. + + _Mus cinnamoneus_ Lichtenstein, 1830, Darstellung neuer oder wenig + bekannter Säugethiere, Berlin, Heft 7, pl. 36. + + _Echimys myosuros_ Is. Geoffroy Saint-Hilaire, 1838, Comptes + Rendus Acad. Sci., 6(26):886, and 1840, Mag. Zool., Paris (ser. 2, + année 2):15, 33, 53; Allen, 1899, Bull. Amer. Mus. Nat. Hist., + 12(20):261. + + _Echinomys leptosoma_ Wagner, 1843, Schreber's Säugethiere, + suppl., 3:341. + + _Echinomys myosuros_ Burmeister, 1854, Syst. ubersicht Thiere + Brasiliens, p. 199. + + _Proechimys setosus_ Thomas, 1921, Ann. Mag. Nat. Hist., 8 (ser. + 9):141. + + _Proechimys myosuros_ 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. + + _Proechimys leptosoma_ 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. + + _General characters._--Aristiforms wide (1''') and numerous + on dorsal parts of body; tail longer (9'') than head and body + (8''); hind feet short (1''6'''). + + _Color_ (According to Lichtenstein and Brants' + descriptions).--Black between ears; dark brown on middorsal line + with reddish tinge on front and upper side of neck; posteriorly + from shoulders there is a greasy shine added to color; this dark, + dorsal band widens posteriorly, there encroaching on sides of + body; sides lighter brown, sparsely marked with dark brown lines, + from sides of head caudad to, and including, outer surfaces of + hind legs; outer sides of forelegs colored like outer sides of + hind legs; ankles ringed with brown. Tail blackish above, whitish + below. Upper surfaces of hands and feet white. + + _Skull._--No description of skull or teeth found. + +_Remarks._--Thomas (1921:140-143) summarized the available information +on the forms from southeastern Brazil and synonymised _myosuros_ with +_setosus_. The description of _leptosoma_ by Brants applies to this same +species except in a few features. However, neither Brants nor +Lichtenstein described the tail of _myosuros_ as having a white tip or +even as having a heavily pencilled tip, although Wagner (1843:342) in +redescribing Lichtenstein's species indicated that the tail had a white +pencil. He gave also measurements of the head and body, and tail, which +do not agree with the original measurements given by either Lichtenstein +or Brants. Lichtenstein, in the original description, gave the type +locality of _myosuros_ as Bahia, and stated that it was collected by +Freireiss. Brants also gave Bahia as the type locality for _leptosoma_. +The names _leptosoma_ Brants and _cinnamoneus_ Lichtenstein are +evidently no more than duplicate names for _myosuros_, as pointed out by +Thomas (1921:141) and subsequent writers. Specimens referred to any of +these forms, therefore, could take the earlier name _myosuros_ until +identified otherwise. Lichtenstein later (1830, text for plate 36, fig. +2) added São Paulo State to the known range of the species, mentioning +specimens collected there by Sello. Probably, therefore, Wagner +redescribed _leptosoma_ using a composite sample; the white tip on the +tail could occur in any race of _P. iheringi_ from São Paulo. + +Considering the short hind feet and the wide aristiforms, _Proechimys +myosuros_ probably will eventually prove to be related to _albispinus_; +perhaps it will prove to be a synonym of _albispinus_. + + + + +CONCLUSIONS + + + 1. The genus _Proechimys_ is divisible into two subgenera. In all + Brazil there are four full species of each subgenus, or 8 species + in all. All but one of these are divisible into subspecies of which + there are 29, making a total of 30 kinds in Brazil; 14 of these are + here newly named. + + 2. It is new information, I think, that: (1) One main fold + extending entirely across the worn crown of the molariform tooth + is peculiar to _Trinomys_; in the subgenus _Proechimys_, apparent + complete division of the crown surface is accomplished by a short + main fold meeting a counterfold originating on the opposite side + of the tooth; (2) progressive decrease in size of molariform teeth + from P4 to M3 is peculiar to the subgenus _Trinomys_; in the + subgenus _Proechimys_, M2 is largest and the teeth are + progressively smaller anteriorly. + + 3. In the one species, _Proechimys albispinus_, which has the + widest distribution of aristiforms on the body of any species in + the genus, some of the aristiforms are clavate. Clavate + aristiforms occur in the most spiny species of the related genus + _Echimys_. + + 4. In subspecies of any one full species the incisive foramen is + larger in animals which inhabit arid areas than in those which + inhabit humid areas. Possibly increased area of moist mucosa + associated with Jacobson's organ is required in arid areas for + maintenance of the necessary keenness of smell. + + 5. The number of counterfolds in the molariform teeth vary in + clinal fashion. Their variation is in response to humidity. + Increasing humidity is correlated with increasing number of folds, + and decreasing humidity is correlated with decreasing number of + folds. + + 6. Clinal variation correlated with increasing humidity is to be + seen also in longer tail and darker color of pelage. + + 7. The primitive _Proechimys_ probably was large, with a short + tail, narrow aristiforms, strongly built skull, and five + counterfolds in each molariform tooth. + + 8. Geographic isolation appears to have been a factor in the + establishment of the two subgenera; the arid belt along the São + Francisco River and northward to Ceará appears to be uninhabited + by _Proechimys_ and constitutes a barrier separating the two + subgenera, _Proechimys_ and _Trinomys_. + + 9. This arid belt probably developed relatively early, since in + deposits of late Pleistocene age, remains of the subgenus + _Trinomys_ have been found in the area where the subgenus still + occurs. + + 10. The most primitive types occur at the periphery of the range + of the genus. + + 11. Populations from small islands tend to be more primitive than + populations on the mainland. Insular populations develop a + homozygous condition with resultant disappearance of secondary + biotypes. + + 12. Insular animals ordinarily are larger than their mainland + counterpart. + + + + +TABLE OF MEASUREMENTS + +TABLE 1.--Measurements (in millimeters) of adults of _Proechimys_ + + Key: + + A Length of head and body + B Length of tail + C Length of hind-foot + D Length of ear from notch + E Greatest length of skull + F Condylo-incisive length + G Zygomatic breadth + H Length of nasals + I Interorbital constriction + J Palatilar length + K Crown length of cheekteeth + + ====================================================================================== + A B C D E F G H I J K + -------------------------------------------------------------------------------------- + _P. g. steerei_, [M] [M] Hyutanaham + USNM 105535 218 123 48 17 53.5 44.0 25.2 19.3 11.7 18.2 8.2 + USNM 105536 217 135 50 55.2 45.3 25.7 20.7 11.4 19.2 8.0 + ? + USNM 105537 56.3 44.9 24.0 20.4 11.4 19.3 8.7 + -------------------------------------------------------------------------------------- + _P. g. goeldii_, [M] Fazenda Paraiso + AMNH 37489 218 52 20 55.1 44.9 27.0 22.1 12.0 18.6 9.4 + [F] + AMNH 37488 228 157 49 22 57.3 47.6 27.9 22.1 12.4 20.3 9.6 + ====================================================================================== + _P. s. liminalis_, [M] [M] Rio Quichito + Mean 229 145 43 21 58.4 47.6 27.3 21.8 12.9 20.1 8.3 + Maximum 250 45 24 61.7 50.0 28.9 23.0 13.8 21.2 8.8 + Minimum 210 40 18 53.3 44.3 25.5 19.4 12.0 18.5 7.9 + No. of specimens 5 1 5 5 5 5 5 5 5 5 5 + [F] [F] + MN 6253 215 150 43 20 57.5 46.5 28.3 21.5 12.0 19.4 8.7 + MN 6250 213 42 20 60.8 49.7 22.0 13.7 21.0 9.1 + -------------------------------------------------------------------------------------- + _P. s. amphichoricus_, [M] [M] Esmeralda + + AMNH 77000 252 163 53 60.8 50.3 27.5 24.4 13.7 20.7 9.4 + AMNH 76994 260 160 54 25.5 22.8 12.6 9.5 + AMNH 77020 250 181 57 62.0 52.0 27.3 25.9 13.8 21.5 9.6 + [F] + AMNH 76999 235 149 50 24.0 19.0 9.3 + -------------------------------------------------------------------------------------- + _P. s. kermiti_, [F] Lower Solimões + AMNH 37124 210 55 65.2 53.7 29.2 27.6 13.5 21.4 9.0 + -------------------------------------------------------------------------------------- + _P. l. brevicauda_, [M] João Pessoa + DZ 900 245 147 48 58.3 46.6 26.5 22.5 11.6 18.2 8.2 + -------------------------------------------------------------------------------------- + _P. l. boimensis_, [M] Boim + MCZ 30881 220 160 50 54.6 44.2 24.8 21.0 11.7 17.3 7.5 + [F] + MN 1976 182 140 45 52.6 42.2 24.3 20.8 11.4 16.8 7.6 + [F] Cametá + MCZ 30878 240 48 55.1 45.0 25.2 22.4 11.5 18.4 7.6 + -------------------------------------------------------------------------------------- + _P. l. longicaudatus_, [M] Urucum + CNHM 26732 229 121 48 21.5 11.5 18.5 8.3 + [F] [F] + AMNH 37085 210 150 44 51.4 42.5 24.3 19.5 10.8 17.1 8.9 + AMNH 37086 210 50 48.5 40.9 23.7 17.1 10.2 17.1 8.3 + -------------------------------------------------------------------------------------- + _P. l. leucomystax_, [M] Tapirapoã + AMNH 37509 230 150 43 50.7 42.2 24.5 18.4 10.3 17.5 8.1 + [F] + AMNH 37510 210 42 48.1 41.3 23.0 16.9 9.9 17.2 8.1 + [F] Utiarití + MN 2212 48.0 39.9 18.3 11.2 15.9 8.0 + [F] Salto Sepotuba + MN 1936 202 147 44 52.6 43.0 23.6 19.2 10.7 17.8 7.8 + -------------------------------------------------------------------------------------- + _P. l. roberti_, [M] [M] Anapolis + Mean 208 159 45 21 52.7 43.5 24.9 20.7 13.1 17.2 7.9 + Maximum 235 190 55 25 56.1 47.8 27.1 23.7 12.0 19.1 8.2 + Minimum 170 135 36 18 48.1 40.0 22.8 18.2 10.6 15.6 7.6 + No. of specimens 16 14 16 16 11 11 11 11 11 11 11 + [F] [F] + Mean 219 149 44 20 51.1 42.3 24.1 20.0 10.7 17.2 8.0 + Maximum 290 155 48 24 55.5 45.4 25.8 21.5 11.1 17.7 8.7 + Minimum 195 125 40 18 48.9 40.3 23.1 19.1 10.5 16.6 7.7 + No. of specimens 10 8 10 10 7 7 7 7 7 7 7 + ====================================================================================== + _P. g. villicauda_, [M] [M] Tapirapoã + MN 1932 225 145 47 55.6 45.5 26.8 24.0 12.0 18.1 8.9 + MN 1934 215 162 50 56.2 46.0 26.1 21.3 12.0 18.6 8.4 + [M] Utiarití + AMNH 57544 250 200 55 . 24.3 13.1 19.9 9.1 + -------------------------------------------------------------------------------------- + _P. g. ribeiroi_, [M] Rio 12 de Outubro + MN 1935 190 134 47 50.1 41.0 24.3 20.0 11.5 15.9 8.1 + -------------------------------------------------------------------------------------- + _P. g. hyleae_, [M] [M] Tauarí + Mean 248 146 52 58.1 47.6 27.1 22.9 12.1 19.4 8.5 + Maximum 260 174 53 59.0 47.8 . 23.4 13.2 20.0 8.8 + Minimum 217 143 51 57.2 46.5 . 22.4 11.1 18.8 8.3 + No. of specimens 4 3 4 2 2 1 2 2 2 2 + [F] [F] + Mean 229 149 50 54.3 44.9 25.8 21.0 11.8 18.1 8.5 + Maximum 270 168 54 56.1 46.3 27.4 23.0 13.4 19.3 9.0 + Minimum 190 132 49 51.5 42.9 24.5 19.1 11.1 17.1 7.9 + No. of specimens 10 9 10 9 11 11 10 10 10 10 + -------------------------------------------------------------------------------------- + _P. g. nesiotes_, [M] Ilha de Manapirí + CNHM 19496 201 133 47 20 52.7 42.7 25.1 19.5 11.1 17.8 8.0 + [F] + MN 1975 200 152 47 21 52.1 42.6 25.8 19.5 12.3 18.3 8.0 + -------------------------------------------------------------------------------------- + _P. g. leioprimna_, [F] [F] Cametá + AMNH 37484 192 151 41 22 54.8 44.9 25.4 19.2 12.6 18.4 8.4 + CNHM 19503 189 164 47 22 54.4 43.7 26.2 20.5 12.6 18.2 8.2 + CNHM 19536 189 43 22 + -------------------------------------------------------------------------------------- + _P. g. oris_, [M] Providencia + CNHM 19495 230 170 45 24 56.1 47.1 25.8 22.0 11.4 17.4 8.3 + [M] Tanaquará + MN 1974 230 175 49 23 53.2 43.0 23.8 20.1 10.5 17.9 7.7 + [M] Rio Guamá + AMNH 37487 205 142 42 21 + -------------------------------------------------------------------------------------- + _P. g. arescens_, [M] [M] Fazenda Inhuma + CNHM 26440 206 149 51 24 54.7 44.1 26.3 21.4 12.1 18.7 8.3 + CNHM 26441 191 164 51 55.6 45.0 25.7 22.4 11.7 18.7 8.7 + -------------------------------------------------------------------------------------- + _P. g. riparum_, [F] Manaus + AMNH 143018 225 44 20 52.6 43.2 24.0 20.5 11.0 17.2 6.7 + -------------------------------------------------------------------------------------- + _P. g. arabupu_, [M] [M] Arabupu + AMNH 75816 243 220 56 59.2 48.7 27.0 23.8 12.9 17.8 8.7 + AMNH 75819 230 181 52 55.0 46.0 26.0 22.5 12.3 16.9 8.3 + AMNH 75815 228 198 52 + [F] [F] + AMNH 75810 226 170 48 53.9 45.6 25.6 21.0 12.0 16.1 8.3 + AMNH 75823 209 188 48 53.4 43.9 24.5 21.5 11.7 16.4 8.3 + AMNH 75817 204 167 47 51.1 43.1 . 21.6 11.3 16.6 8.2 + -------------------------------------------------------------------------------------- + _P. dimidiatus_, [M] [M] Pedra Branca + Mean 199 170 46 52.4 43.5 26.2 19.5 12.1 16.4 8.3 + Maximum 220 195 50 56.4 47.1 27.5 21.5 13.6 18.0 8.7 + Minimum 180 150 44 48.1 40.4 24.6 17.5 11.0 14.4 7.4 + No. of specimens 19 18 19 45 46 45 45 46 46 46 + [F] [F] + Mean 197 162 44 51.8 42.9 25.8 19.4 11.8 16.3 8.3 + Maximum 230 180 46 55.1 45.9 27.4 22.0 13.0 18.4 8.9 + Minimum 165 145 42 48.6 39.5 23.8 17.6 10.7 14.8 7.7 + No. of specimens 14 12 14 42 44 44 42 44 44 44 + -------------------------------------------------------------------------------------- + _P. i. iheringi_, [M] [M] Ilha de São Sebastião + Mean 207 197 48 54.5 44.6 26.2 19.9 12.0 18.3 8.3 + Maximum 220 205 50 55.0 45.2 27.1 20.4 12.8 18.7 8.5 + Minimum 196 190 46 53.5 43.7 25.9 19.3 10.9 17.5 8.0 + No. of specimens 5 2 5 5 5 4 4 5 5 5 + [F] [F] + MN 6453 228 185 46 54.3 44.5 25.9 20.5 11.0 18.9 8.2 + DZ 2095 205 180 46 53.2 42.9 26.4 18.9 11.1 17.0 8.2 + DZ 2525 205 46 56.9 45.8 27.8 20.9 12.5 18.9 8.2 + -------------------------------------------------------------------------------------- + _P. i. bonafidei_, [M] [M] Fazenda Bõa Fé + Mean 211 186 50 25 53.3 43.4 25.6 20.2 12.1 16.3 8.5 + Maximum 220 194 54 26 55.8 45.2 26.3 21.7 13.2 17.4 9.1 + Minimum 200 176 47 24 50.7 41.0 24.2 19.1 11.0 14.8 8.1 + No. of specimens 7 5 8 8 4 5 5 7 7 7 7 + [F] [F] + Mean 209 185 52 25 52.6 44.4 26.6 20.0 12.4 16.4 8.6 + Maximum 226 203 55 27 56.9 46.4 28.6 21.4 13.2 17.0 9.2 + Minimum 185 153 50 22 44.4 42.3 24.9 18.0 11.4 15.5 8.2 + No. of specimens 7 7 7 7 6 6 7 7 7 7 7 + -------------------------------------------------------------------------------------- + _P. i. gratiosus_, [M] [M] Floresta da Caixa Dagua + Mean 193 191 48 51.2 41.4 25.5 18.5 11.7 16.3 8.0 + Maximum 200 200 49 51.7 42.2 27.0 18.9 12.0 16.7 8.2 + Minimum 185 175 47 50.5 40.4 24.6 18.0 11.1 15.6 7.9 + No. of specimens 5 5 5 5 5 5 5 5 5 5 + [F] [F] + Mean 204 175 49 24 50.5 42.1 26.0 18.4 11.4 16.3 7.9 + Maximum 220 190 50 26 52.6 43.1 26.6 19.5 12.1 17.5 8.2 + Minimum 195 160 47 22 48.4 41.0 25.3 17.5 10.7 15.3 7.6 + No. of specimens 5 2 5 4 4 4 4 4 4 4 4 + -------------------------------------------------------------------------------------- + _P. i. paratus_, [M] [M] Floresta da Capela de São Braz + MN 4023 203 195 54 28 51.2 41.7 25.4 18.3 10.4 16.2 8.4 + MN 5458 190 170 51 27 + [F] + MN 4012 220 210 54 29 52.2 42.3 25.4 19.1 12.3 17.5 8.7 + -------------------------------------------------------------------------------------- + _P. i. panema_, [M] [M] Campinho + MN 8286 215 45 23 54.0 45.1 27.7 19.5 13.4 16.4 8.1 + MN 8284 195 180 43 23 51.5 41.8 24.5 18.1 11.4 16.0 7.6 + [F] [F] + MN 8288 190 190 46 21 51.6 42.8 25.3 18.1 11.7 15.7 7.9 + MN 8287 200 190 46 23 52.6 43.6 27.2 18.4 12.5 16.7 8.3 + MN 8285 190 45 24 50.0 41.1 26.6 19.4 12.3 16.1 8.1 + -------------------------------------------------------------------------------------- + _P. i. denigratus_, [M] [M] Mata do Ribeirão da Fortuna + Mean 197 218 52 24 51.5 42.2 25.7 18.3 11.3 16.0 8.2 + Maximum 217 242 54 28 55.4 45.3 27.0 20.3 12.4 17.6 8.5 + Minimum 190 204 50 21 48.7 39.5 23.7 16.8 10.4 15.0 8.0 + No. of specimens 10 9 9 10 8 8 8 8 8 8 8 + [F] [F] + Mean 201 207 52 24 49.1 41.2 25.0 18.3 11.2 16.0 7.8 + Maximum 225 225 54 28 54.1 44.6 25.7 21.5 11.8 17.1 8.3 + Minimum 180 175 49 20 48.2 39.5 23.5 17.4 10.5 15.5 7.5 + No. of specimens 12 12 12 12 8 8 8 8 8 8 7 + -------------------------------------------------------------------------------------- + _P. s. setosus_, ? No locality + AMNH 16140 39.8 25.2 11.0 15.9 7.6 + -------------------------------------------------------------------------------------- + _P. s. elegans_, [M] Lagoa Santa + UZM H82 190 190 47 25 24.3 17.5 11.2 16.6 7.7 + ? + UZM L104 48.6 40.8 24.8 16.6 10.3 15.9 7.7 + ====================================================================================== + _P. a. albispinus_, [M] [M] Macaco Seco + Mean 175 162 40 45.9 39.7 23.6 15.6 10.6 15.7 7.6 + Maximum 193 173 47 48.2 42.0 24.9 17.1 11.6 17.5 8.2 + Minimum 165 147 38 44.2 38.1 22.7 14.8 9.7 14.3 7.2 + No. of specimens 9 8 9 11 12 12 14 13 14 13 + [F] [F] + CNHM 20402 187 171 40 46.0 40.8 23.8 14.9 11.3 16.0 7.4 + CNHM 20408 47.6 40.7 24.6 17.1 11.2 15.9 8.2 + -------------------------------------------------------------------------------------- + _P. a. sertonius_, [M] Lamarão + CNHM 18196 190 160 36 25 11.7 + [M] Bonfim + MN 6454 185 175 37 45.7 39.8 24.2 15.5 10.7 16.1 7.5 + [F] [F] + DZ 2636 190 150 35 43.8 37.6 22.8 14.7 9.6 14.5 + DZ 2635 175 155 38 46.7 40.6 23.3 16.9 10.2 15.1 7.4 + -------------------------------------------------------------------------------------- + + + + +LITERATURE CITED + + + ALLEN, J. A. + 1904. Mammals from the District of Santa Marta, Colombia, collected + by Mr. Herbert H. Smith, with field notes by Mr. Smith. Bull. Amer. + Mus. Nat. Hist., 20:407-468, November 28, 1904. + + 1916. Mammals collected on the Roosevelt Brazilian Expedition, + with field notes by Leo E. Miller. Bull. Amer. Mus. Nat. Hist., + 35:559-610, August 9, 1916. + + AMEGHINO, F. + 1934. Notas sobre una pequéna coleccion de huesos de mamiferos, + procedentes de las grutas calcáreas de Iporanga en el Estado São + Paulo (Brasil), Obras completas y Correspondencia Cientifica de + Florentino Ameghino, La Plata, 17:103-153. + + BERRY, E. W. + 1942. Mesozoic and Cenozoic plants of South America, Central + America and the Antilles. Proc. Eighth Amer. Sci. Congress, + 4:365-373. + + DESMAREST, A. G. + 1817. Nouveau Dictionaire d'Histoire Naturelle, ed. 2, 10:59. + + ELLERMAN, J. R. + 1940. The families and genera of living Rodents. Vol. 1, Rodents + other than Muridae, pp. xxvi + 689, 189 text figs. British Museum + (Natural History). June 8, 1940. + + GEOFFROY SAINT-HILAIRE, ISIDORE + 1840. Notice sur les rongeurs épineux, désignés par lés auteurs + sous les noms d'_Echimys_, _Loncheres Heteromys_ et _Nelomys_. Mag. + Zool. (Guerin-Meneville), s. 2, Ann. 2, pp. 1-57, pls. 20-29. + + LICHTENSTEIN, M. H. C. + + 1830 (1827-34). Darstellung neuer oder wenig bekannter Säugethiere, + 65 Arten, etc., pp. 118, 50 pls. colored. + + LUND, P. W. + 1841. Blik paa Brasiliens Dyreverden för sidste Jordomvaeltning, af + Dr. Lund. Kongelige Danske Videnskabernes Selskabs + naturvidenska-belige og mathematiske. Fortsaettelse af + Pattedyrene. Kjöbenhavn, 2:217-272, pls. 14-24. + + OLIVEIRA, A. I., and LEONARDOS, O. H. + + 1943. Geologia do Brasil. Min. da Agricultura. Rio de Janeiro, 26 + + 813, 202 figs., photographs, 1 map. + + OSGOOD, WILFRED H. + 1944. Nine new South American rodents. Zool. Ser. Field Mus. Nat. + Hist., 29:191-204, July 12, 1944. + + RIBEIRO, ALIPIO DE MIRANDA + 1914. Historia Natural. Zoologia. Mammiferos. Commissão de Linhas + Telegraficas, Estrategicas de Matto Grosso ao Amazonas. Annexo no. + 5, Rio de Janeiro, pp. 1-49 + 1-3, pls. 25, May, 1914. + + RIDGWAY, ROBERT + 1912. Color standards and color nomenclature, iv + 44 pp., 53 pls. + Published by the author, Washington, D. C. + + TATE, G. H. H. + 1935. The taxonomy of the genera of Neotropical hystricoid rodents. + Bull. Amer. Mus. Nat. Hist., 68:295-447, June 12, 1935. + + 1939. The mammals of the Guiana Region. Bull. Amer. Mus. Nat. + Hist., 76:151-229, October 20, 1939. + + THOMAS, OLDFIELD + 1900. Descriptions of new rodents from western South America. Ann. + and Mag. Nat. Hist., 6(ser. 7):294-302, September, 1900. + + 1905. New Neotropical _Molossus_, _Conepatus_, _Nectomys_, + _Proechimys_, and _Agouti_, with a note on the genus _Mesomys_. + Ann. and Mag. Nat. Hist., 15(ser. 7):584-591, June, 1905. + + 1912. On small mammals from the Lower Amazon. Ann. and Mag. Nat. + Hist., 9(ser. 8):84-90, January, 1912. + + 1920. On mammals from the Lower Amazons in the Goeldi Museum, + Pará. Ann. and Mag. Nat. Hist., 6(ser. 9):266-283, September, + 1920. + + 1921. On the spiny rats of the _Proechimys_ group from + Southeastern Brazil. Ann. and Mag. Nat. Hist., 8(ser. 9):140-143, + July, 1921. + + WAGNER, ANDREAS + 1843. Beschreibung einiger neuer Nager, welche auf der Reise des + Herrn Hofrats v. Schubert gesammelt wurden, mit Bezugnahme auf + einige andere verwandte Formen. Abhandl. Akad. Wiss. math.-phys. + Cl., 3(4):173-216, pl. 2. + + WAGNER, J. A. + 1844 (1774-1846). In Schreber's Die Säugethiere ..., 7 pts. + including text and pls., colored (347). + + WIED, MAXIMILIAN ZU + 1826. Beiträge zur Naturgeschichte von Brazilien, vol. 2, Mammalia, + 600 pp., 6 pls., Weimar. + + WINGE, HERLUF + 1888. Jordfundne og nulevende Gnavere (Rodentia) fra Lagoa Santa, + Minas Gerais, Brasilien. E Museo Lundii Afhandlinger, 1(3):1-178, + pls. 1-8. + + 1941. The interrelationships of the mammalian genera. Translated + from Danish by E. Deichmarm and G. M. Allen. København, 3 vols. + (vol. 1, xii + 418), 3 pls., 1941; vol. 2, 376 pp., 1941; vol. 3, + 308 pp., 1942. + + + _Transmitted, December 1, 1947._ + + + 22-3343 + + + + +Transcriber's notes: + +- inconsistent use of m (for meter/s) throughout the whole book. + e.g.: 100 m or 120 m, etc. +- 'TABLE 1.--Measurements (in millimeters) of adults of _Proechimys_' + left in its (remaining) width ( > col. 75) for best possible overview. + + + + + +End of the Project Gutenberg EBook of Speciation in the Brazilian Spiny Rats, by +João Moojen + +*** END OF THE PROJECT GUTENBERG EBOOK 42720 *** |
