Speciation in the Brazilian Spiny Rats

Part 1

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

Part 2

CONCLUSIONS 393

TABLE OF MEASUREMENTS 395

LITERATURE CITED 400

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.

Part 3

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.

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