HOLKY UBRARY KBo 71995 UNlvtRSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY ||g Q r Contributors to Fieldian; a Field Mui.eum staff members anc sidered as space permits N1:i'u:srr:D! all four ov \< jitor. Mitific Eiditor, /'■ iced o ible cornput - ubmit a "Table of Contei title page. In most cases, the text should be pre nd "Literature Cited." io system (periods are not used after abbre 'iait journal titles sho ttlv should follow papers) or S, mica! or TL ,-rsity Press, Stanford, Calif J., J. R. Lloyd, and T. D. Penning V comparison of montane an rest structure, j •. I I ultural patterns in visions, pp. 63-i 'tars. Mouton Publishers, The Hague, ••■p. 785-821. In S H., ed., Ha >f Guatemala. Part II. Polypodiaceae. Fieldiana: i :s" in the text (not as "plates"). Figi narked on the re gure captions ent issues of Fieldiana (< name, figure number(s), and "top." dm .n the arrangement to be obtained i ;and inkdr ...■'• ■ . ipon pu b I espondmg author nade and queries he single set id changes in page JfON IS PRINTED ON ACiD-FREE PAPER. ^ u X2 § c o o M a u ^. 0 3 0 X FIELDIANA Zoology NEW SERIES, NO. 75 A New Central Brazilian Genus and Species of Sigmodontine Rodent (Sigmodontinae) Transitional between Akodonts and Oryzomyines, with a Discussion of Muroid Molar Morphology and Evolution Philip Hershkovitz Curator Emeritus Division of Mammals Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496 Accepted April 16, 1993 Published September 30, 1993 Publication 1453 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1993 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 Material and Methods 1 Classification 2 mlcroakodontomys, new genus (superfamily muroidea) 2 mlcroakodontomys trans1torius, New Species 2 Description 3 Comparisons 4 Systematic Affinity 4 Fringe Zone Faunas 5 Pace of Sigmodontine Evolution 10 Molar Crown Patterns: A Review 10 The Mesoloph (id): Evolutionary Significance 11 Acknowledgments 17 Literature Cited 18 Left side of skulls shown in Figure 1 , with corresponding mandibles 6 Upper and lower right molars of speci- mens shown in Figure 1 7 Diagram of occlusal surface of an upper first molar 12 Diagram of evolution from complex to simple of the occlusal surface pattern of an upper second muroid molar 13 Some enamel patterns of tetralophodont molars 14 Transition of the enamel pattern of an upper second molar from pentalophodont to tetralophodont 15 Diagram of occlusal surface of a lower right first molar 16 List of Tables List of Illustrations 1 . Skulls, dorsal and ventral aspects, of Mi- croakodontomys transitorius, Oligoryzo- mys sp., Oligoryzomys ni gripes, Microry- zomys minutus Measurements of Microakodontomys transitorius and comparable sigmodon- tine rodents 8-9 Frequency of mesolophostylid in molars of Oligoryzomys sp 10 A New Central Brazilian Genus and Species of Sigmodontine Rodent (Sigmodontinae) Transitional between Akodonts and Oryzomyines, with a Discussion of Muroid Molar Morphology and Evolution Philip Hershkovitz Abstract Monotypic Microakodontomys, a long-tailed sigmodontine rodent, inhabits the fringe zone between open grassland and bordering scrubland of the Brazilian cerrado. It most nearly re- sembles the oryzomyine Oligoryzomys but differs by pointed, blackish striped muzzle, cranial proportions, absence of stapedial and sphenofrontal foramina, and loss of mesoloph (id) in all molars. Whereas the derived dental character is an adaptation for a harsh pastoral diet, as in vole-like akodontines, the primitive oryzomyine-like long tail and feet are adaptations for climbing and better suited for sylvan life. The character mosaic marks a transition from the sylvan to pastoral habitus within an ecological fringe zone. The generic description is followed by a discussion of muroid molar crown morphology and its evolution from the primitive pentalophodont pattern characteristic of sylvan mice, to the derived tetralophodont pattern characteristic of pastoral mice, and the phylogenetic implications. Introduction The heretofore undescribed long-tailed mouse discovered in the Brazilian cerrado resembles small species of the widespread sigmodontine Oligory- zomys. Its molars, however, each with mesoloph (id) absent, are unlike those of Oligoryzomys or any oryzomyine. In contrast, sigmodontines with mesoloph-less, brachyodont, tuberculate molars are, as a rule, short-tailed, terrestrial herbivores of open or scrub country. The sylvan-pastoral phenotype parallels that of certain other oryzomyine-thomasomyine-like sig- modontines such as Pseudoryzomys and Wiedo- mys with reduced or absent mesolophs (ids) and habitat in a fringe zone between retreating forests and advancing savannas. Those mice also appear to be transitional between their hypothetical syl- van ancestral morphs characterized by complex pentalophodont molars and their ultimate derived pastoral morphs with simplified tetralophodont molars. Material and Methods The holotype of the new form was taken in a snap trap, its skull damaged with zygomata and right half of braincase lost during hand cleaning and its prepared study skin in poor condition. Di- rect comparisons were made with 18 individuals of an undescribed species of Oligoryzomys taken in the same fringe zone, 4 in the same trapline. Also used in comparison were representatives of other small sigmodontines, including Oligoryzo- FIELDIANA: ZOOLOGY, N.S., NO. 75, SEPTEMBER 30, 1993, PP. 1-18 mys nigripes, of the same region and elsewhere, and the two known species of Andean Microry- zomys. The material collected in Brasilia, D.F., in 1986 by the author and associates of the Field Museum of Natural History (fmnh), are registered in the Museu Nacional, Rio de Janeiro (mnr). Compar- ative material, collected in 1988 in Iporanga, Sao Paulo, by the same personnel, are entered in the registry of the Museu de Zoologia, Universidade de Sao Paulo (uspmz). Cranial measurements (in mm) and terminology for evolutionary stages of cricetid molar crown topography are described elsewhere (Hershkovitz, 1990b, pp. 27, 33). Classification South American mice are here regarded as form- ing a monophyletic group of the subfamily Sig- modontinae. Unresolved is the family to which the subfamily belongs. Some assign the taxon to the Muridae, others to the Cricetidae. In any case, it is generally accepted that these and all other families of mice and rats constitute the superfam- ily Muroidea. In their arrangement of the major components of the Muroidea, Carleton and Musser (1984, p. 294) placed the Sigmodontinae, Cricetinae, Mu- rinae, and others with the Muridae, but with some equivocation. They asked, "Should rats and mice, be assigned to just one family, to the two families Muridae and Cricetidae, or to more than two?" Their answer, " We don't know" (p. 299). The "ar- rangement of the groups as equivalent subfamilies of Muridae," they explained, "reflects not our con- viction that this is the preferred nomenclature, but rather our uncertainty of the hierarchical pattern and our design to focus upon the distinctive and richly varied geographic and biological properties of the groups comprising the Muroidea." The following description of a new genus with a new species focuses on the environmental factors and processes involved, primarily those related to the designs of the molars, in sigmodontine radi- ation. As for the family designation, it is generally held that the "Cricetidae" appeared in upper Eo- cene, the "Muridae" much later, in Middle Mio- cene. There is no consensus, however, regarding lineal relationship, and none regarding origin of South American sigmodontines. Treatment of the group as simply muroid will serve for the nonce. Microakodontomys, New Genus (Superfamily Muroidea) Genotype— Microakodontomys transitorius, new species. Diagnosis— One of smallest sigmodontines with most external and cranial characters as in com- parably small, long-tailed, pentalophodont ory- zomyines but all molars without mesoloph (tetra- lophodont), snout pointed, each side of rostrum with blackish stripe, stapedial and sphenofrontal foramina absent. Remarks— The genus combines the tetralopho- dont molar crown pattern characteristic of pas- toral rodents such as akodonts and phyllotines with most external and cranial characters of such pen- talophodont sylvan rodents as oryzomyines. Over- all resemblance to Microryzomys and Oligoryzo- mys suggests differentiation from an oryzomyine stock in a scrub brush habitat transitional between forest and savanna. The following detailed description and com- parisons combine those of the genus with its ge- notype and only species. Microakodon tomys transitorius, New Species Holotype (frontispiece)— Young adult male, skin and skull, mnr, no. 25986, collected 14 July 1986, by Philip Hershkovitz and Scott Morrow Lindbergh, original no. ph 9515. Skull with right half of braincase, zygomatic arches, and posterior half of each mandible missing. Type Locality— Parque Nacional de Brasilia, D.F., about 20 km NW of Brasilia, D.F. (15°47'S, 47°55'W); taken on edge of campo limpo (wet val- ley side grassland), cerrado biome, at about 1 100 m altitude where Akodon lindberghi, Thalpomy cerradensis, and Oligoryzomys sp. near flavescens were captured during the same period. Distribution— Known from the type locality only. Judged by the single capture during 5 weeks of intensive trapping, the species is probably rare. The Parque Nacional had been continuously FIELDIANA: ZOOLOGY trapped for small mammals during the last quarter century by other investigators, mainly ecologists, without indication in their reports that another specimen of Microakodontomys had been taken or observed. Description External (frontispiece)— Size small; pelage soft, somewhat lax, upper parts ochraceous-buff lined with blackish, more saturate on head, the cover hairs dark brown basally, broadly banded buffy subterminally, narrowly tipped blackish; project- ing guard hairs blackish; eyes rimmed blackish; muzzle with narrow blackish band on each side from tip of snout to corner of eye; sides of trunk paler than dorsum, the guard hairs shorter and fewer; underparts and inner surface of limbs washed with ochraceous; tail longer (133%) than com- bined head and body length, dark above, terminal two-thirds beneath paler; mystacial vibrissae short, the longest extending back to ears; hindfeet long, narrowly palmate basally, inunguiculate pedal dig- it I extending to base of phalanx of digit II, digits II and IV nearly as long as longest digit III, digit V extending to base of phalanx 3 of digit IV; entire plantar surface coarsely scutulated, the interdigital and hypothenar plantar tubercles small, padded thenar tubercle elongate; claws thin, moderately long, not concealed by sparse whitish digital vi- brissae, the manual slightly longer than the pedal, the middle claw longest. Cranial (figs. 1-2)— Rostrum moderately long, slender; nasals about 34% greatest skull length, tapered with pointed tip not projecting beyond incisors; facial lamina of lachrymal bone slightly inflated; interorbital region narrow, smooth, slightly constricted medially, comparatively little divergent posteriorly, fragmented zygomatic arch- es with only anterior upright plates preserved, the flaring right suggesting widespread nearly parallel- sided zygomata; zygomatic plate slightly project- ing, anteroposterior diameter of superior notch of infraorbital foramen about 0.87 mm, or length of m1; capsule of incisor root partially overlapped by anterior margin of zygomatic plate; dorsal contour of braincase flat above, moderately curved behind; interparietal bone long, wide; inflated portion of supraoccipital bone visible dorsally; incisive fo- ramina narrowly ovate and extending to anterior margin of first molars; palatal bridge produced beyond plane of last molars the border with large posterolateral pits; mesopterygoid fossa at anterior base of pterygoids wider than either paraptergoid fossa at same transverse plane; length of auditory bulla slightly greater than length of upper molar row; stapedial and sphenofrontal foramina absent, squamosal-alisphenoid groove absent. Fragmentary mandible (fig. 2) with angular notch deep, angular process extending posteriad to pos- terior rim of condylar process; incisor root capsule little projecting. Dental (fig. 3)— Upper incisors opisthodont, the outer enamel orange; upper molars brachyodont, tuberculate; outer and inner upper cusps opposing, the outer slightly higher than inner; mesoloph (id) absent in all molars. (Numbers in parentheses are of elements shown in figs. 4-8.) M1— Anterocone with well-defined anterolabial and anterolingual conules, anteromedian style not distinct or fused with anterolabial conule; antero- labial conule, anterolophule, plesiostyle, and para- style fused isolating paraflexus (3) from margin; paralophule, metalophule, and ill-defined meso- style fused isolating metaflexus (5) from margin; anterior fossette absent; median fossette opening into mesoflexus (4); posterior fossette coalesced with metaflexus; low entoloph extending to margin fused with entostyle in left molar, absent in right; posteroflexus (6) obsolete; posterostyle and pos- teroloph absent; protoflexus (8) and hypoflexus ( 1 0) broad, deep; enterostyle present; distoflexus not indicated. M2 — Paraflexus (3) isolated from margin by fu- sion of mesolophule with anteroloph and para- style; median fossette discrete; posterior fossette indicated; posteroflexus (6) present; mesostyle present; enterostyle present; short protoflexus (8) present; hypoflexus (10) broad, deep; enterostyle present. M1— Outline subtriangular, less than half bulk m2, cusps low; isolated paraflexus (3) and meta- flexus (5) extremely short; median fossette greatly enlarged; protoflexus (8) weakly defined; hypo- flexus (10) short, compressed. Lower molars subhypsodont, terraced, the inner cusps crested, the outer plane, cusps of each side alternating; protoflexid (8) well defined in all 3 molars, hypoflexid (10) broad, deep, median fos- sette confluent with mesoflexid (4), posterior fos- sette confluent with posteroflexid (6). M, — Well-defined anterolingual and anterola- bial conulids subequal; anterolophid absent, an- terostylid fused with metaconid isolating antero- HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT flexid (2) from margin; posterolophulid absent; mesostylid and posterostylid present, each isolat- ing mesoflexid (4) and posteroflexid (6), respec- tively, from margin; median and posterior fos- settes each coalesced with corresponding mesoflexid and posteroflexid; ectolophid and sty- lids between outer cusps absent. M2 — Mesoflexid and posteroflexid isolated; me- sostylid well developed; each fossettid probably coalesced with corresponding flexid; ectolophid absent, ectostylid present; hypoflexid (10) broad, deep. M3— Subtrapezoidal, about two-thirds bulk of m2; hypoconid flat like protoconid; lingual flexids isolated. Comparisons (figs. 1-3) The comparisons apply to the genus and lone species of Microakodontomys. Microakodontomys transitorius is hardly distin- guishable externally from Microryzomys minutus Tomes or Microryzomys altissimus Osgood except for smaller ears and blackish rostral stripes. It also resembles the slightly larger syntopic Oligoryzo- mys sp. with ears the same size but rostral stripes absent. A series of the latter was taken in the same habitat. The larger Oligoryzomys nigripes was taken during the same trapping period in other nearby more humid localities. Cranially the holotype is grossly similar to Mi- croryzomys spp. and Oligoryzomys spp. except di- astema shorter, rostrum narrower, interorbital re- gion more nearly parallel-sided, less divergent behind, braincase narrower, the parietals not glo- bose, nasals tapered proximally, the tips bluntly pointed, not rounded and not overhanging the in- cisors. The partially damaged left half of braincase (right half missing) resembles that of Oligoryzomys spp. by absence of sphenofrontal foramen and as- sociated squamosal-alisphenoid groove. It differs from those of Microryzomys spp. and most Oli- goryzomys spp. by absence of stapedial foramen. Dentally, complete absence of mesoloph (id) in all molars separates M. transitorius from oryzo- myines, thomasomyines, and all other pentaloph- odont rodents and from all tetralophodont rodents with mesoloph short or vestigial. The tetralophodont molar pattern combined with the cranial characters may seem to align the holotype with akodont rodents, particularly the smaller species of Akodon including the syntopic A. lindberghi (Hershkovitz, 1990a, p. 16). It is distinguished from like-sized akodonts, however, by one or more characters such as tail longer than length of head and body combined, hindfeet long and narrow, claws unspecialized, upper molars brachyodont, tuberculate, interorbital region smooth and parallel-sided with little divergence behind, incisive foramina short, bullae small. Morphometric comparisons with small species of Akodon appear in Table 1. Additional measure- ments and characterizations of larger akodonts and related forms are given by Hershkovitz (1990b, p. 4) and by Myers et al. ( 1 990). Compared with like- sized phyllotines such as Calomys spp., tail con- siderably longer than combined head and body length, hindfeet longer, zygomatic plate less pro- jecting, interorbital region narrower, mesoptery- goid fossa wider incisive foramina shorter. In no way could Microakodontomys be derived from any phyllotine with molars like those of Calomys (sen- su stricto) or its molariform parallel, the North American Blancan Bensonomys Gazin. For char- acters and dimensions of Calomys, see Hershko- vitz (1962, pp. 124, 189). The blackish rostral stripes of Microakodonto- mys transitorius recalls facial patterns of the long- clawed sigmodontine Geoxus valdivianus and many didelphoids (Marsupialia). Oxymycterus para- mensis nigrifrons also shows the blackish midros- tral stripe with extensions to outer corner of each eye in a few individuals. Summary— Judged only by the fragmented skull and damaged skin, M. transitorius could be taken for a well-differentiated species of Oligoryzomys most nearly related to O. flavescens, or O. fornesi, or the larger O. nigripes. Absence of mesoloph (id) in all molars, blackish rostral bands, and tail length more than 1 1 0% of combined head and body length, however, distinguishes Microakodontomys transitorius not only from oryzomyines but from all other sigmodontines of comparable or other body size. Systematic Affinity Microakodontomys, like eastern Brazilian Wie- domys, Pseudoryzomys, and Rhagomys, retains the long tail, long foot, scansorial proclivities, and gen- eral appearance of its postulated sylvan oryzo- myine-thomasomyine origin. No intimate rela- tionship, however, is inferred between any of the genera. Each appears to be too widely divergent FIELDIANA: ZOOLOGY Fig. 1 . Skulls, dorsal and ventral aspects (registry no. and greatest length in mm in parentheses); white bar = 1 cm. A, Microakodontomys transitorius (mnr 25969; 21.5); B, Oligoryzomys sp. (mnr 25949; 22.3); C, Oligoryzomys nigripes, young (mnr 27083; 23.3); D, Microryzomys minutus (fmnh 71907; 20.5). dentally and cranially for alignment with other genera or generic groups derived from either the oryzomyine or thomasomyine branch. In his assessment of the Sigmodontinae, Reig (1980, p. 263) disposed of the eastern Brazilian Rhagomys and Pseudoryzomys, the Peruvian al- tiplano Punomys, and the intercontinental low- land Zygodontomys as "Sigmodontinae incertae sedis." As for Wiedomys, Reig (1980, p. 265) com- bined it with his newly described Colomys from the Argentine Pleistocene to form the new tribe Wiedomyini. Other sigmodontine tribes recog- nized were the Recent Oryzomyini, Akodontini, Phyllotini, Sigmodontini, Scapteromyini, and Ichthyomyini. To these may be added the dis- tinctive Thomasomyini and Oxymycterini. Pend- ing further study, Microakodontomys, perhaps an independent offshoot of an oryzomyine branch of the presumed monophyletic sigmodontine tree, is tentatively cast as Sigmodontinae incertae sedis. Fringe Zone Faunas Microakodontomys transitorius is phenotypi- cally and functionally transitional between sylvan and pastoral sigmodontines. Its habitat is the ce- rrado fringe or broad, intergrading zone between sylvan and pastoral environments. The bordering forests are isolated Amazonian outliers at the headwaters of the Rio Tocantins. Mice of these gallery forests are sylvan. Those of the cerrado are pastoral. Fringe faunas, on the other hand, include sylvan, pastoral, and transitional species. Most HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT Fig. 2. Left side of skulls shown in Figure 1 , with corresponding mandibles; white bar (size of d slightly exaggerated) = 1 cm. abundant and adaptive fringe zone mice are the wide-ranging species of Oligoryzomys. The cerrado grades from wet campo limpo or grassland, to campo sujo or mixed grassland, shrub, and low, widely scattered trees. Campo sujo, in turn, merges into campo cerrado with its thick cover of brush, bushes, and low trees. Microako- dontomys transitorius was taken in the fringe be- tween campo limpo and campo sujo in the same trapline as a large series of a superficially similar species of Oligoryzomys and the larger, short-tailed Akodon lindberghi Hershkovitz (1 990b) and Thal- pomys cerradensis Hershkovitz (1990a, pp. 763 et seq. for habitats and sigmodontine fauna). The tetralophodont molar pattern of M. transi- torius is primarily adapted for mastication and Fig. 3. Upper and lower right molars of specimens shown in Figure 1 (greatest length of tooth rows in mm in parentheses). A, Microakodontomys transitorius (3.1/3.2); B, Oligoryzomys sp. (3.1/3.2); C, Oligoryzomys nigripes (3.6/3.8); D, Microryzomys minutus (3.1/3.2). FIELDIANA: ZOOLOGY Table 1 . External and cranial measurements (in mm) of Microakodontomys transitorius and some like-sized Sigmodontinae (Oryzomyines and Akodonts). Head and body Tail : head and Hind foot Taxon Locality length Tail length body length ratio length with claw Microakodontomys Brazil: transitorius Brasilia 70 93 133 21 Microryzomys Peru altissimus* 74.6(62-84)22 108(95-122)22 144f 21(18-23)22 Oligoryzomys Brazil: sp. <5<5 Brasilia 79(70-86)10 105(97-111)10 133(117-149)10 22(21-23)10 Oligoryzomys Brazil: sp. 22 Brasilia 82(75-87)5 102(95-111)5 125(114-137)5 21.6(21-22)5 Oligoryzomys Brazil: nigripes 66 Iporanga 96(83-105)16 122(113-146)16 127(107-145)16 24(23-25)16 Oligoryzomys Brazil: nigripes 22 Iporanga 99(93-107)5 125(113-129)5 125(119-130)5 24(21-26)5 Akodon lutescens§ Peru: Arequipa 78(76-81)5 60(57-64)5 76(70-84)5 19.4(19-20)5 Akodon caenosus§ Argentina: Jujuy 98(85-103)9 60(56-64)9 61(55-66)9 19(18.5-19.5)9 Akodon dolichonyx Chile: Atacama 87(77-94)14 55(47-66)14 63(52-78)9 20.3(19-22)14 Akodon alterus§ Argentina: Catamarca 91,89, 100 63,61,78 69, 68, 78 19, 19,20 Akodon puer% Bolivia: Cochabamba 84(75-94)5 71(70-75)5 85(80-93)5 18(15-21)5 Akodon boliviensis% Peru: Puno 99(90-115)11 62(55-66)11 63(52-72)11 21(20-22)11 * Data from Carleton and Musser (1989, p. 81) except as noted with f. t Mean tail : mean head and body length, and other measurements. % fmnh, Peru (La Quinua). § According to Myers et al. (1990, p. 62), A. lutescens and A. caenosus are each subspecies of A. puer; A. pacificus is a synonym of A. boliviensis, whereas A. spegazzini is distinct from tucumanensis, a synonym. Table 1. Continued. Nasals, Rostral Rostral Zygomatic Taxon Locality length length width plate Microakodontomys Brazil: transitorius Brasilia 7.7 6.4 3.6 1.9 Microryzomys Peru altissimus* 6.3(5.5-7.6)5f 6.35(5.5-7.6)17 3.9(3.7^.0)5 1.8(1.7-2.0)5t Oligoryzomys Brazil: sp. 66 Brasilia 8.4(8.0-9.2)10 7.9(7.6-8.2)10 4.1(3.9^.3)10 2.4(2.0-2.7)10 Oligoryzomys Brazil: sp. 29 Brasilia 8.4(7.8-8.9)5 7.6(7.1-8.1)5 3.9(3.8^.0)5 2.2(2.1-2.3)5 Oligoryzomys Brazil: nigripes 66 Iporanga 9.4(8.5-10.4)15 8.8(7.8-9.2)13 4.2(3.7-4.9)14 2.4(2.2-2.6)16 Oligoryzomys Brazil: nigripes 22 Iporanga 9.4(9.0-9.9)5 8.6(9.1-9.0)5 4.3(4.1-4.7)5 2.4(2.2-2.6)5 Akodon lutescens§ Peru: Arequipa 7.8(7.3-8.1)4 8.2,8.2 4.1(3.9-4.2)5 1.9(1.6-2.1)5 Akodon caenosus§ Argentina: Jujuy 8.5(7.7-8.5)9 7.8(7.3-8.4)9 4.2(4.0-4.3)9 2.0(1.9-2.2)9 Akodon dolichonyx Chile: Atacama 8.7(7.6-9.3)9 7.9(7.5-8.3)5 4.2(3.5^.5)9 2.2(2.0-2.3)9 Akodon alterus§ Argentina: Catamarca 8.5,8.3,9.2 8.9, 8.4, 9.5 4.2, 4.4, 4.6 2.2, 2.2, 2.3 Akodon puer§ Bolivia: Cochabamba 8.9,8.1,8.4 8.2,8.3,8.8 4.0(3.7^.1)4 2.2(2.0-2.5)4 Akodon boliviensis§ Peru: Puno 8.5(7.9-9.8)9 8.8(8.5-10.0)11 4.6(4.4-4.8)10 2.2(2.0-2.6)10 FIELDIANA: ZOOLOGY Table 1. Continued. Ear length from notch Greatest skull length Zygomatic breadth Interorbital width Frontoparietal suture Braincase width 13 22.8 — 2.9 6.7 9.2(= 4.6 x 2) 15.3(15-16)5* 22.2(20.0-24.9)13 11.3(10.7-12.4)15 3.3(3.0-3.6)22 8.1(7.5-8.7)5t 10.8(10.2-11.2)18 13(13-14)10 23.0(22.3-23.3)10 12.2(11.9-12.7)10 3.6(3.4-3.7)10 7.9(7.4-8.4)10 10.9(10.4-11.5)10 14(13-15)5 22.8(22.4-23.3)5 12.0(11.6-12.6)5 3.6(3.4-3.8)5 7.5(7.2-7.7)5 10.9(10.1-11.3)5 16(15-17)16 24.9(24.1-26.3)15 12.9(11.9-13.6)16 3.4(3.2-3.6)16 7.2(6.9-7.8)15 11.2(10.6-11.6)16 16.6(16-18)5 25.1(24.2-25.9)5 13.0(12.5-13.4)5 3.4(3.1-3.6)5 7.3(7.1-7.5)5 11.0(10.9-11.2)5 22.3(22.0-22.7)4 11.1(10.7-11.3)5 4.1(4.0-4.2)5 7.7(7.0-8.2)5 10.7(10.4-10.9)5 12(12-12)5 23.6(23.0-23.9)9 11.7(11.5-12.1)9 4.3(4.1^4.4)9 7.8(7.3-8.4)9 10.9(10.4-11.2)9 23.4(22.4-24.4)8 11.9(11.5-12.6)8 4.0(3.8^4.1)8 7.8(7.2-8.3)5 11.3(10.9-11.8)8 13, 13, 13 24.0, 22.8, 26.0 12.2. 12.0, 12.4 4.2,4.4,4.3 7.6, 7.4, 7.9 -, 11.0, 11.6 15(15-15)4 24.4, 22.9, 23.7 11.7(11.4-12.3)4 4.3(4.2^4.6)4 7.9(7.6-8.5)4 10.8(10.5-11.2)4 14.5(14-15)10 24.6(23.8-25.6)9 12.6(11.8-13.2)8 4.2(4.1^4.4)10 7.7(6.9-8.3)9 9.2(8.6-10.0)10 Table 1. Continued. Incisive Palatal M" M" M' Diastema foramina bridge length across width 4.7 3.5 3.8 3.1 3.8 0.86 5.4(4.8-6.2)31 4.3(3.5-4.6)20 3.5(3.1^4.0)23 3.1(2.8-3.3)21 4.0(3.4-4.4)21 0.9(0.8-1.0)25 5.5(5.2-5.8)10 4.4(4.0-5.1)10 4.0(3.7-4.2)10 3.2(3.1-3.4)10 4.1(4.0-4.4)10 1.0(0.86-1.2)10 5.3(5.2-5.3)5 4.5(4.1-4.7)5 3.8(3.7^4.0)4 3.14(3.1-3.2)5 4.1(4.0-4.3)5 1.0(0.91-1.1)5 5.9(5.5-6.2)15 5.1(4.5-5.8)16 4.1(3.6-4.6)16 3.6(3.2-3.9)16 4.2(3.9-4.6)15 1.01(0.86-1.27)15 5.8(5.7-6.0)5 5.2(4.9-5.3)5 4.0(3.7^4.3)5 3.6(3.3-3.7)5 4.3(4.0-4.6)5 0.9(0.8-1)5 5.5(5.4-5.7)5 5.4(5.1-5.7)5 3.1(3.0-3.2)5 3.6(3.4-3.7)5 4.5(4.3-4.6)5 0.98(0.94-1.0)5 6.0(5.6-6.2)9 5.7(5.1-6.1)9 3.1(2.7-3.4)9 3.5(3.4-3.7)9 4.7(4.5-5.0)9 0.98(0.92-1.0)9 5.3(4.8-5.6)5 5.2(4.7-5.7)10 3.5(3.2-3.6)10 3.4(2.8-3.7)9 4.7(4.5-5.0)5 1.1(1.0-1.2)5 6.0, 5.9, 6.5 5.7,5.5,6.4 3.0, 3.3, 3.6 4.3, 4.0, 4.4 4.8,4.8,4.7 1.1, 1.1, 1.3 5.8(5.5-6.3)4 5.7(5.4-6.2)4 2.9(2.8-3.1)4 3.6(3.5-3.7)4 4.8(4.5-5.1)4 1.0(0.94-1.1)4 6.4(5.7-7.2)10 5.6(5.2-5.9)10 3.3(3.0-3.8)10 4.0(3.9^4.1)10 5.0(4.8-5.3)10 1.1(1.0-1.3)11 HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT Table 2. Frequency of fused mesolophid and me- sostylid (= mesolophostylid) in lower molars of Oligo- ryzomys sp. (N = 18) from Parque Nacional, Brasilia D.F.; R = right, L = left. Fused mesoloph and mesostyle (= mesolophostyle) is present in all upper molars. Present Absent M R L R L M, M, M3 Totals 10 5 16 31 10 7 16 33 8 8 13 11 2 2 23 21 secondarily for grinding harsh forage. The long tail and moderately long feet are suited both for ter- restrial and scansorial locomotion and for gaining nesting sites above ground level. In open grass- land, a long tail on a small mouse could be dis- advantageous. Molar differentiation ofMicroakodontomys must have occurred during a long dry period when erst- while extensive humid forests shrank to moist banks of streams or into tracts isolated by sur- rounding savannas. A syntopic species in transit from pentalophodonty to tetralophodonty is the undescribed fringe zone Oligoryzomys sp. men- tioned above. All 18 specimens captured in the same trapline as the holotype of M. transitorius retain the sylvan externalities including long tail, long foot, and general oryzomyine appearance as in M. transitorius. Upper molars of the series are entirely pentalophodont, the mesolophostyle in- tact in each tooth. Lower molars, however, are about equally pentalophodont and tetralopho- dont. Those most frequently tetralophodont are the second lower molars, at the center of masti- catory impact. Least tetralophodont are the third lower molars, the smallest and least functional of the suite judged by wear. Patterns ofm,, largest of the molars, are mixed (table 2). Pace of Sigmodontine Evolution Emergence of the semipastoral Microakodon- tomys with a clear stamp of oryzomyine origin, and discovery of the syntopic Oligoryzomys sp. in partial transition to the pastoral habitus, suggests rapid speciation, perhaps less than a millenial pace. The fossil record indicates, however, that many living sigmodontines had already attained present generic and perhaps species grade, no more re- cently than late Pliocene or early Pleistocene. The common sigmodontine ancestral form, likely near living thomasomyines, remains to be discovered, perhaps in South America, possibly in Eurasia or Africa. Molar Crown Patterns: A Review (figs. 4-8) The morphology, terminology, and functional and ecological significance of pentalophodont and derived tetralophodont molar crown patterns, ba- sic to an understanding of sigmodontine and other rodent radiations, have been described by Hersh- kovitz (1944, 1955, 1960, 1962). They are sum- marized here. The ancestral complex or pentalophodont cri- cetid molar is characterized by five primary trans- verse crests (Hershkovitz, 1962, p. 76). Those of the upper molars are I, procingulum; II, paracone; III, mesoloph fused with mesostyle, or mesoloph- ostyle; IV, metacone; and V, postcingulum. Those of the lower molars are I, procingulid; II, meta- conid; III, mesolophid fused with mesostylid, or mesolophostylid; IV, entoconid; and V, postciijgu- lid. All molars of oryzomyine, thomasomyine, and the North American peromyscine Ochroto- mys, Megadontomys, Aporodon, and Isthomomys are pentalophodont. Mice with pentalophodont molars are primarily sylvan. In most deforested environments, sylvan forms may find refuge in gallery forests, wooded ravines, shrub-covered hillsides, banks of brush-lined streams, and the fringe zones between strictly sylvan and pastoral habitats. The derived or simplified tetralophodont molar is marked by the same primary transverse crests except that crest III, the primitively fused meso- loph-mesostyle (id), is disjunct, with either or both elements reduced or absent. Sometimes the me- sostyle (id) alone becomes hypertrophied in ab- sence of the mesoloph (id). The evolutionary process of molar crown sim- plification, beginning with the breakup of crest III, proceeds with reduction and elimination of trans- verse crests IV and I (the order sometimes re- versed), followed by merger of II and IV (Hersh- kovitz, 1962, fig. on p. 77, pp. 82-101). Modifications commence with the lower molars, the second usually earliest. Mice with tetralopho- dont molars are all New and Old World living muroids not mentioned above. Old World mu- 10 FIELDIANA: ZOOLOGY rines lack even a vestige of the mesoloph. Tetra- lophodont sigmodontines are primarily pastoral; their habitats include deserts, savannas such as tundras, punas, scrublands, cerrados, and caatin- gas, and palm, bamboo, and coniferous forests. These mice may also occur as intrusive, isolated, or fringe elements in relict or refuge bogs, mead- ows, rocky situations, and partially cleared or cul- tivated fields and along grassy or brush-lined banks of streams cutting through broad-leaf forests. The mesoloph is lost in transition from the ancestral sylvan pentalophodonty of sylvan forms to de- rived tetralophodonty of pastoral forms. Revers- ibility of the process, mentioned in a review by Voss (1991, p. 37), is improbable. An element once lost is never regained. The narrow, fluctuating fringe between forest or thick shrub and open grassland may be foraged by rodents natural to either pastoral or sylvan habi- tats or transient between them. Seasonally chang- ing diets in fringe zones, however, include harsh fiber plant foods selective for a tetralophodont mo- lar pattern. Pseudomesolophs (ids)— Paralophules or me- talophules are evaginations of corresponding up- per cusps; metalophulids or entolophulids are equivalent extrusions of corresponding lower cusps. The lophules (ids) may appear singly or in com- bination in one or more of the pentalophodont or tetralophodont molars of a suite. A lophule (id), particularly of the tetralophodont molar, may sim- ulate the mesoloph (id), often by fusion with it, the mesostyle (id), or both. Any one or combi- nation of these protrusions and fusions of lophules (ids) are the pseudomesolophs (ids) described by Hershkovitz (1962, p. 80). The true mesoloph (id) is a lateral protrusion or evagination of the mid- sagittal spine or mure (id) of the molar and no other part of the tooth. It may fuse partially or completely with any or all enamel elements be- tween paracone and metacone (metaconid and en- toconid) as well as with the cones (ids) themselves (fig. 7B). Failure to recognize the distinction between true mesoloph (id) as an evagination or development of the mure only, and its relationship with the mesostyle (id), on the one hand, and the various lophules (ids), on the other, obscures phylogenetic relationships between species, genera, and higher categories of sigmodontines. Vorontsov ( 1 979) and Hooper (1957) adopted the molar enamel pattern introduced by Hershkovitz ( 1 944, 1 955) but never fully comprehended the taxonomic, much less phylogenetic, significance of the true mesoloph (id) or mesolophostyle (id). Others before and since have confused this element with pseudomesoloph (id) variables of little or no significance in the tran- sition from pentalophodonty to tetralophodonty (cf. Lindsay & Jacobs, 1985, p. 24, fig. 8e, pis. 2ij). Hooper (1957, p. 9, fig. 1 on p. 14) provided an excellent description of the mesoloph (id), but the exposition exceeds the limitations of the subject. In his figure, the true mesoloph (id) is shown pro- truding directly and correctly from the mure in upper and lower second molars. The inset enlarge- ments of various secondary positions of purported mesolophs (ids), however, are misleading. His in- set figures 1-4, and possibly 5 of mJ are paraloph- ules; the same numbered figures of m2 are ento- lophulids. Not shown are the equivalent metalophules and metalophulids. All may be cat- egorized as pseudomesolophs (ids), but the correct term for the single element is lophule (id). The true mesoloph (id; fig. 7B) may fuse with any or all lophules (ids) or with the principal cones (ids) themselves. The Mesoloph (id): Evolutionary Significance The passage from pentalophodonty to tetra- lophodonty with reduced to absent mesoloph is as significant in muroid evolution as is the transition from trituberculy to quadrituberculy in earliest ro- dents, primates, and other mammalian orders. It compares with the gradual loss of digits in ungulate evolution from woodland browser to pastoral grazer. The eight sigmodontine dental characters enu- merated by Jacobs and Lindsay (1984, p. 269, table 2) as primitive include no. "4 short mesoloph (and mesolophid)." Molars with reduced or absent mesolophs (ids) are derived. They are almost the only kinds preserved as fossil in former pastoral or fringe habitats. Their characters were the ones considered by Jacobs and Lindsay (1984, p. 269) in their construction of sigmodontine phylogeny. Their putative ancestor, the middle Miocene Megacricelodon with mesoloph of m ' reduced, as in many living and extant muroids, was already too derived to have given rise to the Sigmodon- tinae, contra Jacobs and Lindsay (1984, p. 269, fig. lb). On the other hand, Jacobs and Lindsay (1984, p. 269) were correct in rejecting Leidymys as a likely ancestor of sigmodontines. They were wrong in attributing this postulate of ancestry to Hershkovitz. HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT 11 LABIAL LINGUAL RIGHT M1 Fig. 4. Diagram of occlusal surface of a first right upper molar showing basic elements of the enamel pattern in muroid rodents. 12 FIELDIANA: ZOOLOGY Fig. 5. Diagram showing evolution from complex to simple of the occlusal surface of a generalized upper second muroid molar: A, complex or pentalophodont molar pattern with the five transverse buccal crests I-V (lingual in lower molar); B, simplified or tetralophodont pattern initiated by disjunction of mesoloph and mesostyle; C, tetraloph- odont molar pattern with loph III reduced or lost, and reduction, obsolescence, or elimination of lophs I and V; D- F, further simplification of the tetralophodont pattern from bilophodont (D-E) to cylindroform (F). For explanation of symbols see below. For b' read a', for c' read b'. Explanation for symbols of Figures 4-7 (a-i in- clusive = procingulum or loph I; s, t, z = postcin- gulum of loph V); the fold terminology (1-11) is from Hershkovitz (1962, p. 70). a. anteromedian style (may be fused with b, c, or both) b. anterolabial lophule c. anterolingual lophule d. anterolophule (may be fused with 0 e. anterior fossette f. plesiostyle (may be fused with d, h, or j) g. protostyle (may be fused with i) h. anteroloph (may be fused with f, j, or both) i. protoloph (may be fused with g, u, or both) j. parastyle (may be fused with f, h, k. or a combination) k. mesolophule (may be fused with h, j, or both) 1. paracone m. paralophule (may be fused with n, o, or both); element may be multiplied n. mesoloph (when fused with o = mesolopho- style) o. mesostyle (may be fused with m, p, or both; when fused with n = mesolophostyle) p. metalophule (may be fused with o, n, or both) q. metacone r. posterolophule (may be fused with s) s. posterostyle (may be fused with r, t, or both) t. posteroloph t'. posteroconule (usually not differentiated from posteroloph, t) u. protolophostyle (may be fused with i) v. protocone w. enteroloph (may be fused with x) (continued) HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT 13 IV IV C D E Fig. 6. Some enamel patterns of the tetralophodont upper and lower molars formed by lamination and involution: A, crown pattern formed by lamination (3-8; 5-10), and involution (6), of right m2 of the South American caviomorph Dinomys brannichi Peters; B, pattern by lamination (5-10), and involution (3, 6), of right m2 of the South American caviomorph Dactylomys dactylinus Desmarest; C, formation of an epsilon pattern by involution (3, 5, 10) of left m2 of the South American sigmodontine Phyllotis micropus; D, formation of modified S or sigmoid pattern by involution (10) and elimination (5) of left m3 in Phyllotis micropus; E, formation of S or sigmoid pattern by involution (10, 3) and elimination (8) of left m2 in the South American sigmodontine Euneomys chinchilloides. For explanation of symbols, see pages 13-14, 17. Figure modified from Hershkovitz (1962, p. 96). x. enterostyle (may be fused with w) y. hypocone z. distostyle a', median fossette (may be coalesced with 3, or unite with 4) b'. posterior fossette (may be coalesced with 5, or unite with 6) c'. protolophule d'. hypolophule e'. mure (border between lingual and labial cusps and lophs 1 . preflexus (anterior median fold) 2. anteroflexus (anterior secondary fold) 3. paraflexus (first primary fold) 4. mesoflexus (first secondary fold) 5. metaflexus (in absence of mesoloph coalesced with 4 or first secondary fold) 6. posteroflexus (second secondary fold) 7. supraflexus (anterior lingual fold; in absence of protoloph coalesced with 8 or first minor fold) 8. proton1 exus (first minor fold) 9. entoflexus (major fold) 10. hypoflexus (in absence of enteroloph coa- lesced with 9 or major fold) 1 1 . distoflexus (second minor fold) 14 FIELDIANA: ZOOLOGY "0 0 "CO JQ 1 c Ey / c c^v x-^ >* EN \ <4— - \ K 1 ° r- a CD c 2 C O o j2 5*1 •El "8 ^ ° o 0 i5 o 1 «f 2-g "£ <■ W u 0-4= O - "> U. C wo ° H? - 5 — O. O O. Oy _0 15^ C < a.s 8 lis C 3 _ S'-sJ O u u c , 5*1 ^ E'-o w„ c si S .£ a HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT 15 LINGUAL LABIAL RIGHT Mj Fig. 8. Diagram of occlusal surface of a first right lower molar showing basic elements of the enamel pattern in muroid rodents. 16 FIELDIANA: ZOOLOGY Explanation for symbols of Figure 8 (a-h inclu- sive = procingulum or loph I; a', 14, 16 = post- cingulum or loph I); terminology (1-11) is from Hershkovitz (1962, p. 70). anteromedian stylid (may be fused with b, c, or both) anterolabial conulid (may be fused with c) anterolingual conulid (may be fused with b) labiolophulid (may be fused with g) anterior fossette anterolophulid (may be fused with h, i, or both) prostylid (may be fused with d) anterolophid (may be fused with f, i, s, or combi- nation) i. anterostylid (may be fused with f, h, s, or combi- nation) j. protoconulid (may be fused with g) <. protoconid 1. paralophulid (may be fused with m, n, or both) i. ectostylid (may be fused with 1, n, o, or combina- tion) l. ectolophid (may be fused with 1, m, o, or combi- nation) hypoconulid (may be fused with m or n) hypoconid posterolophulid (may be fused with r) posterolophid (may be fused with q) posteroconulid (usually not differentiated from pos- terolophid, r) s. mesolophulid (may be fused with h) t. metaconid u. metalophulid (may be fused with v, w, or both) v. mesolophid (when fused with w = mesolophostylid) w. mesostylid (when fused with v = mesolophostylid) x. entolophulid (may be fused with v, w, or both) y. entoconid z. distolophulid (may be fused with a') a', posterostylid b'. median fossetid c'. posterior fossetid d . protolophulid e'. hypolophulid f. murid (zone between lingual and labial lophs and cusp) 1 . preflexid (anterior median fold) 2. anteroflexid (anterior labial fold; in absence of an- terolophid coalesced with 3) 3. metaflexid (first secondary fold) 4. mesoflexid (first primary fold) 5. entoflexid (second secondary fold; in absence of me- solophid, coalesced with 4) 6. posteroflexid (second primary fold) 7. supraflexid (anterior labial fold; in absence of la- biolophulid coalesced with 8) 8. protoflexid (first minor fold) 9. ectoflexid (major fold) 10. hypoflexid (in absence of ectolophid [n] coalesced with 9) 1 1. distoflexid (second minor fold) Acknowledgments Fieldwork in Brazil during 1986 was made pos- sible through the sponsorship of Professor Ulises Caramaschi of the Museu Nacional, Rio de Ja- neiro. Permission for faunal investigations in the National Parks and Reserves was granted by the Instituto Brasileira de Desenvolvimento Florestal of the Ministerio de Agricultura. I am beholden to Christopher Tribe, at the time a graduate as- sistant in the Museu Nacional, for cooperation in the fieldwork, and to then Universidade de Bra- silia graduate student Miguel Marini for hospital- ity and volunteer fieldwork. I am especially grate- ful to Professor Jader Soares Marinho Filho of the Universidade de Brasilia, for loan of traps, field assistance, and use of university facilities. In this and continuing investigations of the small mammals of the Brazilian cerrado and the Mata Atldntica, I am deeply indebted to Field Museum Associate Scott Lindbergh, for unstinted cooper- ation in all aspects of the Brazilian small mammal inventory project. Help in the research and preparation of the manuscript by Associate Barbara E. Brown is gratefully acknowledged. The illustrations are the work of volunteer artist Kathleen Kozol Telfer. The scanning electron photographs of skulls and teeth were taken by Field Museum staff botanist Betty Strack. Prints of these and other negatives used here were made by staff photographers Linda S. Dorman and Diane White. Fieldwork and lab- oratory research were supported by the Field Mu- seum Barbara E. Brown Fund for Mammal Re- search. HERSHKOVITZ: NEW CENTRAL BRAZILIAN GENUS OF SIGMODONTINE RODENT 17 Literature Cited Carleton, M. D., and G. G. Musser. 1984. Muroid rodents, pp. 289-379. In Anderson, S., and J. K. Jones, Jr., eds.. Orders and Families of Recent Mammals of the World. John Wiley & Sons, New York. . 1989. Systematic studies of oryzomyine ro- dents (Muridae, Sigmodontinae): A synopsis of Mi- crorvzomvs. Bulletin of the American Museum of Nat- ural History, 191: 1-83. Hershkovitz, P. 1944. A systematic review of the Neotropical water rats of the genus Nectomys (Crice- tinae). Miscellaneous Publications of the Museum of Zoology, University of Michigan, 58: 1-88. . 1955. South American marsh rats genus Hol- ochilus with a summary of sigmodont rodents. Field- iana: Zoology, 37: 639-673. . 1960. Mammals of northern Colombia. Pre- liminary report no. 8: Arboreal rice rats, a systematic revision of the subgenus Oecomys, genus Oryzomys. Proceedings of the United States National Museum, 110:513-568. . 1962. Evolution of Neotropical rodents (Muri- dae) with special reference to the phyllotine group. Fieldiana: Zoology, 46: 1-524. — . 1990a. The Brazilian rodent genus Thalpomys (Sigmodontinae, Cricetidae), with a description of a new species. Journal of Natural History, London, 24: 763-783. . 1 990b. Mice of the Akodon boliviensis size class (Sigmodontinae, Cricetidae) with the description of two new species from Brazil. Fieldiana: Zoology, 57: 1-35. Hooper, E. T. 1957. Dental patterns in mice of the genus Peromyscus. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 99: 1- 59. Jacobs, L. L., and E. H. Lindsay. 1984. Holarctic radiation of Neogene muroid rodents and origin of South American cricetids. Journal of Vertebrate Pa- leontology, 4: 265-272. Lindsay, E. H., and L. L. Jacobs. 1985. Pliocene small mammal fossils from Chihuahua, Mexico. Paleonto- logia Mexicana, 51: 1-45. Myers, P., J. L. Patten, and M. F. Smith. 1990. A review of the boliviensis group of Akodon (Muridae: Sigmodontinae), with emphasis on Peru and Bolivia. Miscellaneous Publications of the Museum of Zool- ogy, University of Michigan, 177: 1-104. Reig, O. A. 1980. A new fossil genus of South Amer- ican cricetid rodents allied to Wiedomys, with an as- sessment of the Sigmodontinae. Journal of Zoology, London, 192: 257-281. Vorontsov, N. N. 1979. Evolution of the alimentary system in myomorph rodents. Translated from Rus- sian, printed and published by the Indian National Scientific Documentation Centre, New Delhi, for the Smithsonian Institution and the National Science Foundation, Washington, D.C., 346 pp. Voss, R. S. 1991. An introduction to the neotropical muroid rodent genus Zygodontomys. Bulletin of the American Museum of Natural History, 210: 1-1 13. 18 FIELDIANA: ZOOLOGY d at Lake Shore Drii HECKMAN U BINDERY INC. |§ JAN 95 — "^^AANNA^1T2ER