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Reproduction, Growth and Development in Two Contiguously Allopatric Rodent Species, Genus Scotinomys

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Reproduction, Growth and Development in Two Contiguously Allopatric Rodent Species, Genus Scotinomys MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 15 1 Reproduction, Growth and Development in Two Contiguously Allopatric Rodent Species, Genus Scotinomys by Emmet T. Hooper & Michael D. Carleton Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN August 27, 1976 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN FRANCIS C. EVANS. EDITOR The publications of the Museum of Zoology, University of Michigan, consist of two series-the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 19 13, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number of pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mailing list for the series. The Miscellaneous Publications, which include papers on field and museum techniques, monographic studies, and other contributions not within the scope of the Occasional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zoology, Ann Arbor, Michigan 48109. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 15 1 Reproduction, Growth and Development in Two Contiguously Allopatric Rodent Species, Genus Scotinomys by Emmet T. Hooper & Michael D. Carleton Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN August 27, 1976 CONTENTS PAGE INTRODUCTION ........................................ 1 MATERIALS AND METHODS ................................ 2 DISTRIBUTION AND HABITATS .............................. 5 ACTIVITY PATTERNS AND FOOD HABITS ...................... 10 Activity .............................................. 10 FoodHabits ........................................... 13 B1;HAVIOR IN TI-IE LABORATORY ............................ 15 Voice ............................................. 15 Young ......................................... 15 Adult ......................................... 17 Sight and Smell ....................................... 20 Illtcrspecific Behavior ................................... 20 Matings ........................................ 20 Behavior ....................................... 21 REI'RODUCTION ........................................ 22 Seasonality in Breeding ..................................22 Gestation .......................................... 24 Litter Size .......................................... 25 Care of Young ........................................28 POSTNATAL GROWTH AND DEVELOPMENT .................... 31 Selectcd Developmental Events ............................ 31 Condition of the Neonate ............................31 Pigmentation and Pelage ............................ 31 Opening of Eyes and Other Features ..................... 33 Locomotion .................................... 34 Growth Rates ........................................ 35 Sexual Maturity ...................................... 37 DlSCUSSlON ............................................ 40 Evolution of the Interspecific Differences ...................... 40 Contiguous Allopatry and Interspecific Competition .............. 47 LITERATURECITED ...................................... 50 ILLUSTRATIONS FIGURE PAGE 1. Scotinomys teguina vocalizing ............................ 1 2 . Distribution of S . teguina ................................ 5 3 . Geographic ranges of S. teguina and S . xcrampelinus .............. 6 4 . Vertical distribution of Scotinomys on slopes of Volcin IrazG and V . Turrialba ...................................... 8 5 . Activity of Scotinomys in the laboratory ...................... 11 6 . Frequencies of activity bouts of different lengths in Scotinomys ...... 12 7 . Sonagrams of parts of a song of teguina ...................... 18 8 . Numbers of litters produced by month in the laboratory in Scotinomys . 23 9 . Distribution of litters by litter size in Scotinomys ................ 26 10. Litter size with respect to the first four litters in Scotinomys ........ 17 1 1 . Photographs of laboratory raised S . teguina and S . xcrampelinus at birth and at one, two and three weeks ...................... 32 12. Relationship of age at eye opening and litter size in Scotinomys ....... 33 13. Semilogarithmic plots of growth in length and weight in Scotinornys .... 36 14. Semilogarithmic plots of growth in length of body and of tail in Scotinonzys ........................................37 15. Semilogarithmic plots of growth in length of hind foot and of ear in Scotinornys ..................................... 38 16. Gain in weight with rcspect to size of litter in S . teguirza ............ 39 17. Geometric growth in body length in Scotinomys ................. 40 18. Growth in body length in terms of adult size in Scotinomys ......... 41 INTRODUCTION A systematic synopsis of Scotinomys in 1972 (Hooper, 1972) summarized the inform ation on morphology, ecology and distribution of the genus. The known populations were found to represent two species, S. xerampelinus and S. teguina, the latter polytypic. The species were shown to be allopatric, but there was evidence that where their ranges abutted they might be locally sympatric. Relation- ships of the populations in those areas were not understood. The data then available suggested that the two species were insectivorous, diurnal, communicated by high frequency sounds, and that they contrasted in several anatomical and behavioral traits. That systematic synopsis posed questions and set the basis for subsequent research. In areas of contiguous sympatry, are the species occupying different microenvironments? Do they compete, and if so, for what resources? How do interspecific anatomical and behavioral differences (e.g., in aggressiveness, thermoregulation, voice (Fig. 1 ), size of eye, and number of mammary glands), relate to altitudinal segregation of the species, reproductive strategies, and possible inter- specific competition? Information on some of the interspecific differ- ences is detailed elsewhere: temperature regulation (Hill and Hooper, 197 1); orbital region and size of eye (Hooper, 1975); karyotypes and accessory reproductive glands (Carleton, Hooper, and Honacki, 1975); and high frequency vocalization (Anderson and Hooper, ms.). Fig. 1. Scotinomys teguina vocalizing. 2 EMMET T. HOOPER & MICHAEL I). CARLETON The present report is concerned principally with four aspects of the biology of Scotinornys reproduction, postnatal growth and development, behavior and food habits, and distribution and habitats. Secondly, it discusses factors and selection pressures whicli possibly were involved in the evolution of different reproductive regimes, behaviors, and ecologies of the species. We wish to express our sincere appreciation to the following: Jorge Campabadal and Costa Rican personnel of the Organization for Tropical Studies, for their assistance in many ways and their patient endurance; Jose' and Roberto Campabadal, for permitting us to work on their fincas; Jose' Castro, for his gracious hospitality; Laurence C. Stuart, for guidance and companionship in Guatemala; Alfred L. Gardncr, Roy W. and Mercedes McDiarmid, Douglas C. Robinson, Norman J. Scott, Gary Stiles, and Wayne Van Devcnder for informa- tion on possible predators of Scotinornj~s.Philip Myers, for critical comments on the manuscript; Robert C. Farentinos, for ideas and aid in obtaining information on vocalizations and behavior; William C. Stebbins, David Anderson, and Harvey Nye, for assistance and usc of facilities of the Kresge Hearing Research Institute; John T. Ilogg and Lawrencc L. Master for conducting and analyzing behavioral tests; Laura Fabbri, for analyses of food of Scotinoi?zys and associated genera; Mark Orsen and Patricia J. Wynne for the linc drawings; and Luis Martonyi and David Bay, for the photographs. Financial support was provided in part by National Science Foundation grants GB-5801 and GB-35500 to Hooper. MATERIALS AND METHODS The animals used in the laboratory werc obtained in Sher~nan live traps baited with peanut butter and rolled oats, and then shipped by air to facilities in the University of Michigan Museum of Zoology. They are from the following Middle American localities. Scotinoinys teguina Costa Rica: Cartago, Tapanti, 1200 m, May 1969, 1;Volcan IrazG , 2850-2940 m, June 1969, 15; March 197 1, 6; August 1972, 13; Puntarenas, Monte Verde, 1430 In, May 1969, 8. Nicaragua: Matagalpa, 9 km N Matagalpa, Santa Marii, de Ostuma, 1300-1400 m, May 1969, 5. Panama: Chirqui , Rio Chirqui Viejo, 1800 m, May 1969, 1. S. .xerampelirzus. Costa Rica: Cartago, Volciin Irazd and Volcin Turrialba, 2700-3000 in, April 1969, 13; Aug~~st1972, 19. The animals were housed in polycarbonate cages (Maryland Plastics Inc.) of two sizes: 26.7 cIn x 48.3 cm x 15.5 cm for mated pairs and 21 cm x 28 cm x 16 cm for single individuals. White pine shavings covered the cage bottom to a depth of 4 cm, and cotton batting and small metal cans were provided for nesting materials. The lab diet consisted of Purina Cat Chow pellets supplied ad libituin together with a daily feeding, usually in early morning, of approxi- IIEPRODUCTION,
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