View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by DigitalCommons@University of Nebraska University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Papers in Natural Resources Natural Resources, School of September 1992 Morphometrics of the Family Emballonuridae Patricia W. Freeman University of Nebraska-Lincoln, [email protected] Cliff A. Lemen University of Nebraska-Lincoln, [email protected] Follow this and additional works at: https://digitalcommons.unl.edu/natrespapers Part of the Natural Resources and Conservation Commons Freeman, Patricia W. and Lemen, Cliff A., "Morphometrics of the Family Emballonuridae" (1992). Papers in Natural Resources. 19. https://digitalcommons.unl.edu/natrespapers/19 This Article is brought to you for free and open access by the Natural Resources, School of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Natural Resources by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in Contributions to Mammalogy in honor of Karl F. Koopman (T.A. Griffiths and D. Klingener, Eds.), Bulletin of the American Museum of Natural History, No. 206, pp. 54–60. Copyright © 1992 American Museum of Natural History. Used by permission. American Museum of Natural History http://www.amnh.org/ Bulletin of the American Museum of Natural History http://library.amnh.org/pubs/bulletin.html R N\~)H, ISo. 206 : 5'4-*b lqq'L Morphometrics of the Family Emballonuridae PATRICIA W. FREEMAN1 AND CLIFF A. LEMEN2 ABSTRACT Morphometric analysis revealed three distinc- all other emballonurids. Phylogenetic studies also tive groups among the genera of emballonurids. separated Taphozous-Saccolaimus as distinctive Taphozous-Saccolaimus is a group distinctive in but included diclidurids among other New World size and shape, particularly cranially. Diclidurids species. Compared with molossids, emballonurids are distinctive in appendicular characters only, es- are morphometrically quite homogeneous. pecially those in the wing. The third group includes INTRODUCTION Miller (1 907) thoroughly described most recently Barghoorn ( 19 77) examined cranial of the extant families of bats and illustrated morphology of a fossil genus, Vespertiliavus, several of the genera with exquisite line draw- and all Recent genera for their possible phy- ings. His work remains valuable for its clarity logenetic relationships, and Robbins and Sar- and for establishing a description of and qual- ich (19 88) produced a phylogenetic study of itative differences among the morphologies the family using protein electrophoresis and of families and genera of Chiroptera. immunology. We have been interested for some years in the quantitative morphological differences among genera within families and among families (Freeman, 19 8 1; Lemen and Free- Thirty-eight meristic characters (27 cranial man, 1981, 1984). In this morphometric and 1 1 appendicular) were studied on 37 spe- treatment of the family Emballonuridae, the cies of emballonurid bats. These are standard sheath-tailed bats, we examine the quanti- measurements and, for the most part, a sub- tative differences among the species and gen- set of those in Freeman (1981). There are era within the family, compare the morpho- some changes, however, because of structural metric groupings with groupings from two differences between molossids and emballo- phylogenetic studies, and describe the mor- nurids. Because there is no comparable third phometric variation between the families phalanx and cartilaginous tip of digit I11 in Ernballonuridae and Molossidae, the free- emballonurids, the quantity measured for tailed bats. both families is the length from the second Early generic treatments of emballonurids phalanx to the tip of that digit. Postorbital include Troughton's (1 925) revision of the and interorbital breadths in some emballonu- Australasian genera Taphozous and Sacco- rids had to be measured inferior to any su- laimus, Sanborn's (1 937) study of American praorbital bone overhanging those breadths species, and Tate and Archbold's (1939) ex- in order to measure the least constriction. amination of the genus Emballonura. More This was particularly true with the diclidu- ' Curator of Zoology and Associate Professor, Division of Zoology, University of Nebraska State Museum, Lincoln, Nebraska 68588-05 14. * Research Associate, Division of Zoology, University of Nebraska State Museum, Lincoln, Nebraska 68588-05 14. 1991 FREEMAN, LEMEN: EMBALLONURID MORPHOMETRICS 55 rids. The quantity SIZE equals the sum of the opogon, G. Taphozous nudiventris, E. natural logs of greatest skull length (which Taphozous perforatus, I. Taphozous theobal- differs slightly from condylocanine length; di, J. Saccolaimus flaviventris, K. Saccolai- used in Freeman, 1984, 1988), zygomatic mus peli, and L. Saccolaimus saccolaimus. breadth, and height of braincase. SIZE cor- The following measurements were taken relates well with weight of the animal (Free- (descriptions and illustrations are in Free- man, 1988). man, 198 1): Cranial: greatest skull length, Both bivariate and multivariate analyses palatal length, maxillary toothrow, upper were used to assess the data, including prin- molariform row, lacrimal width, interorbital cipal components analysis with and without width, postorbital width (POSTORB), zy- a "shearing" function. The shear method de- gomatic breadth, breadth at mastoids, breadth scribed by Bookstein et al. (1985) is partic- of braincase, height of braincase, height of ularly useful in our analysis because it gen- upper canine, length M3 (M3LENGTH), erates a size factor based on within-genus width M3, width at upper canines, width at comparisons and not the first principal com- upper molars, dentary length, dentary-con- ponent of the entire data set, as used in "size- dylocanine length, condyle to M3 length, out." The distinction between the sizeout ap- lower toothrow, moment arm of temporal, proach and shearing is particularly important moment arm of masseter, height of coronoid, in this data set because the bats in the Ta- dentary thickness, height of condyle above phozous-Saccolaimus group are much larger toothrow, height of lower canine, and length than the majority of Emballonuridae. The of condyle; Appendicular: tibia, forearm, third sizeout approach will tend to define shape metacarpal, third metacarpal first phalanx differences between Taphozous-Saccolaimus (PHAL 1M3), third metacarpal second pha- and other bats as size-related differences. This lanx to tip, fourth metacarpal, fourth meta- would indicate that bats of the Taphozous- carpal first phalanx, fourth metacarpal sec- Saccolaimus group are not different in shape. ond phalanx, fifth metacarpal, fifth Shearing does not use the size differences metacarpal first phalanx (PHAL 1MS), and among groups in its definition of size, and in fifth metacarpal second phalanx. this case Taphozous-Saccolaimus is found to Abbreviations used in the text include PC 1, have considerable shape differences com- principal component one; PC2, principal pared to other Emballonuridae (fig. 1). Fi- component two; PC3, principal component nally, simple regression analyses were per- three; H2, sheared component 2; and H3, formed on each character versus the SIZE sheared component 3. quantity. We examined the following species for this study: a. Balantiopterx io, b. Balantiopterx plicata, c. Centronycteris maximiliani, d. Co- We thank curators of the American Mu- leura afra, e. Cormura brevirostris, f. Dicli- seum of Natural History, the Field Museum durus albus, g. Diclidurus ingens, h. Dicli- of Natural History, and the National Muse- durus isabella, i. Diclidurus scutatus, j. um of Natural History for use of their spec- Emballonura alecto, k. Emballonura atrata, imens. Early data-gathering trips by Freeman 1. Emballonura beccarii, m. Emballonura were supported by the Field Museum. Com- monticola, n. Emballonura nigrescens, o. puter analyses were performed and graphics Emballonura raflrayana, p. Emballonura produced by equipment in the newly estab- semicaudata, q. Emballonura sulcata, r. Per- lished Mary B. Totten Center for Biosyste- opteryx leucoptera, s. Peropteryx kappleri, t. matics Technology located in the University Peropteryx macrotis, u. Peropteryx trinitatis, of Nebraska State Museum. Drs. Thomas A. v. Rhynchonycteris naso, w. Saccopteryx bil- Griffiths and Don E. Wilson kindly reviewed ineata, x. Saccopteryx canescens, y. Saccop- the manuscript and Mark Marcuson, staff art- teryx leptura, A. Taphozous australis, B. ist, assisted with graphics. Finally and most Taphozous hamiltoni, C. Taphozous hilde- importantly the senior author thanks Karl gardeae, D. Taphozous longimanus, E. Ta- Friedrich Koopman (pronounced "Cope- phozous mauritianus, F. Taphozous melan- mun" as would KFK) for serving as one of BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 206 A CRANIAL AND APPENDICULAR B CRANIAL AND APPENDICULAR CRANIAL D APPENDICULAR Fig. 1. The first three principal components of an analysis of the emballonurid data set with cranial and appendicular measurements (A and B). Separate analyses using sheared components were run on cranial measurements alone and on appendicular measurements alone (H2 and H3 in C and D). Species are indicated by letters, which are listed in Methods. her mentors at the American Museum; for who unselfishly shares that knowledge; and patiently wading through new methodology for being a continual source of stimulation