JCBPS; Section B; May.2015–July.2015, Vol. 5, No. 3; 2601-2620. E- ISSN: 2249 –1929
Journal of Chemical, Biological and Physical Sciences
An International Peer Review E-3 Journal of Sciences Available online atwww.jcbsc.org
Section B: Biological Sciences
CODEN (USA): JCBPAT Research Article
Is Mouse Hair Morphology Consistent throughout the Suborders Myomorpha and Castorimorpha?
1Britten Sessions, 2 Wilford M. Hess, 3 Michael Rice, 4 Jared Donaldson, and *5 Brad Carmack
1 Department of Chemical Engineering, Brigham Young University, Provo, UT, US 84602, 2 Department of Plant and Wildlife Sciences, Brigham Young University, 3 Department of Microbiology, Brigham Young University, Provo, UT, 4, 5* Department of Biology, Brigham Young University, Provo, UT, US
Received: 13 April 2015; Revised: 30 April 2015; Accepted: 04 May 2015
Abstract: Surface scale patterns of 18 Utah mouse species from the families Cricetidae, Dipodidae, Heteromyidea, and Muridae were studied using scanning electron microscopy (SEM). Hair width, scale length, patterns, and positions in relation to the longitudinal direction of hair were used to characterize differences and similarities in hair morphology between species within the families studied. Previous mouse hair studies have provided data about distinct mammalian underfur and guard hair characteristics; our results confirm that mouse hair morphology consists of distinct guard hair and underfur which were identified by width size and pattern. Statistical analysis was used to characterize hair morphology for the species studied. Keywords: Hair characteristics, identification chart, hair morphology, mouse, Rodentia, SEM, Utah.
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INTRODUCTION
Mice have long lived with and near humans. Found in essentially every terrestrial habitat, mice have learned to adapt to a variety of conditions and environmental surroundings. Each family has evolved to more specially adapt to its given conditions: Animals in the family Muridae have small hands and strong forelimbs allowing them to jump easier. Animals in the family Dipodidae has long back feet and tail to more easily keep its balance, and the family Heteromidae has pouches to more easily transport food1. Although mouse size tends to vary between 5 and 30 cm, the body length, especially the tail, depends upon the need for greater stability and balance, especially when climbing trees1. Natural selection has played a large role in the preservation of and further diversification of the species of mice. The two rodent suborders Myomorpha and Castorimorpha, which include most of the species of mice, constitute 1,671 described species which is nearly a fourth of all mammalian species. Only the order Chiroptera claims a species diversity of comparable magnitude at 1,116 species2, 3. Due to the length of time in which mice have evolved, large variations and diverse characteristics posit mice as excellent subjects for hair studies with scanning electron microscopy (SEM). Hair samples of species from Myomorpha and Castorimorpha, found in Utah, were selected to determine the variation in hair morphology selected from were studied to determine hair scale patterns, ascertain uniformity among the samples, and compare the results to findings of other mammalian hair studies. Various light, electron microscopy, and image analysis procedures have been used to study mammalian hair 4-22. Many studies have helped to further identify and classify hair structure and morphology 23, 26. However, though mammalian hair research in general has expanded, very little research, including classification, had been done on mouse hair 27, 29. In the last thirty years, transmission electron microscopy (TEM) and SEM analyses of mammalian hair have become popular due to their comparative advantages in magnification and resolution 30. Morphological differences in scale structure and patterns of mammalian hair vary significantly from species to species26, 31, 32. The availability of SEM images and morphological analysis of hair of mouse species will be useful for taxonomic, forensic, and archaeology applications 4, 10, 33-37. Due to controversies in classification of hair of mouse species, SEM analysis could provide a useful tool for further identification of individual species of mice1. The purpose of this study, therefore, was to identify and characterize the hair morphology of the selected 18 species of mice, and compare the results to previously conducted studies.
EXPERIMENTAL
Selected mouse hair samples were secured from 18 specimens in the Monte L. Bean Life Sciences Museum at Brigham Young University. To achieve uniform results of both guard hair and underfur, the hair samples were cut at or near the shoulder 38. Previous studies determine that hair morphology from tanned and untanned specimens appears to be identical 38, and unpublished results3. Following previously described procedures 8, 9, 38, hair samples were submerged in distilled water with a drop of Teepol detergent. The samples were sonicated for three minutes to remove dirt and debris. Samples were then rinsed in distilled water and air dried. For each species specimens were positioned on aluminum stubs mounted with carbon film. Notches were filed into stubs to assist with specimen orientation on stubs. However, for some specimens the small size of the hair made it difficult to carefully orient the specimens. 2602 J. Chem. Bio. Phy. Sci. Sec. B, May 2015 – July 2015; Vol.5, No.3; 2601-2620
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The samples were sputter coated with gold and images were recorded with an FEI XL 30 ESEM FEG (FEI Company, Hillsboro, Ore., USA) as described by Castillo, et al 39. Where possible, hair width was measured at the widest points of hair surfaces. Where curvature was present (Figs. 41-43), hair shafts were also measured at points of maximum width. It has been confirmed that hair height can vary considerably even on one shaft of hair. As such, hair height was averaged by taking several measurements on one hair, repeated on as many hairs as necessary to obtain 30 measurements. Each hair was measured using Image J software (http://rsbweb.nih.gov/ij/).
Figs. 1-6: Scanning electron micrographs of mammalian hair. Family Cricetidae: Brush Mouse (1-3) Cactus Mouse (4-6).
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Figs. 7-12: Scanning electron micrographs of mammalian hair. Family Cricetidae: Canyon Mouse (7-9), Deer Mouse (10-11), Northern Grasshopper Mouse (12).
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Figs. 13-18: Scanning electron micrographs of mammalian hair. Family Cricetidae: Northern Grasshopper Mouse (13-15), Pinyon Mouse (16-18).
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Figs. 19-24: Scanning electron micrographs of mammalian hair. Family Cricetidae: Southern Grasshopper Mouse (19-21). Family Dipodidae: Western Jumping Mouse (22-23). Family Heteromyidae: Dark Kangaroo Mouse (24).
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Figs. 25-30: Scanning electron micrographs of mammalian hair. Family Heteromyidae: Dark Kangaroo Mouse (25-26), Desert Pocket Mouse (27-29), Great Basin Pocket Mouse (30).
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Figs. 31-36: Scanning electron micrographs of mammalian hair. Family Heteromyidae: Great Basin Pocket Mouse (31-32), Little Pocket Mouse (33-34), Long-tailed Pocket Mouse (35-36).
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Figs. 37-42: Scanning electron micrographs of mammalian hair. Family Heteromyidae: Long-tailed Pocket Mouse (37), Olive-Backed Pocket Mouse (38-40), Rock Pocket Mouse (41-42).
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Figs. 43-48: Scanning electron micrographs of mammalian hair. Family Heteromyidae: Rock Pocket Mouse (43), Silky Pocket Mouse (44-46). Family Muridae: House Mouse (47-48).
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Figs. 49-53: Scanning electron micrographs of mammalian hair. Family Muridae: House Mouse (49-50), Western Harvest Mouse (51-53).
Hair morphology was classified following the patterns given by Teerink 26. These included three terms to describe scale position in relation to the longitudinal direction of the hair: transversal, longitudinal, and intermediate. Descriptive scale pattern terms were also after Teerink 26. Three new descriptive scale patterns were observed and designated: elongate broad petal (a cross between Teerink’s elongate and broad petal patterns); joint-cusp (interlocking bone pattern); and jagged elongate (similar to Teerink’s elongate petal pattern but with jagged edges).
RESULTS AND DISCUSSION
For each of the orders and species listed below, SEM was used to visualize morphological characteristics of scale patterns of underfur and guard hair. Comparative descriptions are given which have been summarized in Table-1. SEM was also used to measure guard hair and underfur widths and heights (at least 30 measurements for each species), which are summarized in Tables 2 and 3, respectively. Statistical box plot values for guard hair width and height are included in Figures 54,55 and for underfur width and height in Figures 56,57.
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Table-1: Comparative Description of 18 Specimens Studied.
Guard Hair/ Scale Mean Hair Mean Scale Specimen Scale Pattern(s) Underfur Direction(s) Width (μm) Height (μm) Brush Guard Hair Transversal Broad petal/ regular wave/ unequal 28.6 23.7 hastate/ mosaic Underfur Longitudinal/ Joint cusp/ unequal hastate 12.1 11.5 transversal Canyon Guard Hair Transversal Elongate broad petal/ regular wave 21.9 8.00 Underfur Longitudinal Joint-cusp/ broad petal 12.2 7.70 Cactus Guard Hair Transversal Regular wave/ hastate petal/ 27.1 13.5 Underfur Longitudinal Joint-cusp/ elongate broad 14.6 11.4 Deer Guard Hair Transversal Regular wave/ unequal hastate 23.7 7.50 Underfur Longitudinal Elongate broad petal/ regular wave 13.3 8.10 Pinyon Guard Hair Transversal/ Unequal hastate 27.1 6.50 longitudinal Western Guard Hair Transversal Elongate broad petal/ regular wave 19.0 12.8 Harvest Underfur Intermediate Joint-cusp/ regular wave/ unequal 10.3 10.9 hastate Northern Guard Hair Transversal Regular wave 49.0 10.6 Grasshopper Southern Guard Hair Transversal/ Regular wave 34.7 7.50 Grasshopper longitudinal Dark Guard Hair Transversal Regular wave 35.3 8.10 Kangaroo Little Pocket Guard Hair Transversal/ U Regular wave/ 59.9 8.40 shape Great Basin Guard Hair Transversal regular wave/ unequal hastate/ 19.0 12.7 Pocket Underfur Transversal Regular wave 10.4 9.40 Silky Pocket Guard Hair Transversal Unequal hastate/ regular wave 33.0 8.50 Long-tailed Guard Hair Transversal Regular wave/broad petal 26.2 7.00 Pocket Underfur Longitudinal Unequal hastate/ broad petal 12.0 6.20 Desert Pocket Guard Hair Transversal/U Regular wave 75.9 8.60 Shape Rock Pocket Guard Hair Transversal Regular wave 36.9 7.70 Western Guard Hair Transversal Regular wave/ unequal hastate 23.7 7.80 Jumping Underfur Transversal Regular wave/ unequal hastate 13.4 8.70 House Guard Hair Transversal Regular wave/ unequal hastate 29.5 7.80 Underfur Longitudinal Elongate broad/ joint cusp/ 13.2 13.2 Olive-backed Guard Hair Transversal Elongate broad petal/ regular wave 70.8 6.90 Pocket Underfur Transversal/ Regular wave/ unequal hastate/ 23.3 16.4 longitudinal joint cusp
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Table-2: Statistics of guard hair/underfur widths (based on 30 hair measurement).
Guard Hair/ Sample Q1 Median Q3 Standard Deviation Underfur Brush Guard Hair 23.90 28.23 32.67 4.45 Underfur 10.71 12.04 13.52 5.47 Canyon Guard Hair 18.97 21.10 23.32 1.74 Underfur 11.35 12.39 13.29 2.91 Cactus Guard Hair 23.31 26.85 28.54 7.83 Underfur 13.15 15.11 16.41 7.17 Deer Guard Hair 21.27 22.96 25.96 1.74 Underfur 10.87 12.77 14.85 4.35 Pinyon Guard Hair 26.04 27.07 27.64 1.10 Western Harvest Guard Hair 16.48 17.21 21.30 3.87 Underfur 9.62 10.42 11.01 3.53 Northern Grasshopper Guard Hair 45.00 48.90 52.73 2.87 Southern Grasshopper Guard Hair 32.00 34.08 37.98 1.20 Dark Kangaroo Guard Hair 32.91 36.47 37.77 1.50 Little Pocket Guard Hair 57.50 60.20 65.12 1.68 Great Basin Pocket Guard Hair 17.16 17.79 21.16 4.42 Underfur 9.51 10.32 11.25 2.39 Silky Pocket Guard Hair 29.17 34.69 36.55 1.80 Long-tailed Pocket Guard Hair 21.44 25.94 30.50 1.59 Underfur 10.6 11.40 13.65 4.49 Desert Pocket Guard Hair 68.97 75.08 81.35 1.70 Rock Pocket Guard Hair 34.81 36.91 40.05 1.54 Western Jumping Guard Hair 22.84 23.41 25.60 2.09 Underfur 12.01 13.38 14.51 1.66 House Guard Hair 23.83 29.87 34.66 1.26 Underfur 11.25 13.34 14.54 3.92 Olive-backed Pocket Guard Hair 50.66 73.54 91.67 2.18 Underfur 18.15 23.73 29.13 4.76
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Table-3: Statistics of guard hair/underfur heights (base on 30 hair measurement).
Guard Hair/ Standard Sample Q1 Median Q3 Underfur Deviation Brush Guard Hair 23.90 28.23 32.67 7.53 Underfur 5.81 10.62 17.36 2.20 Canyon Guard Hair 18.97 21.10 23.32 3.85 Underfur 5.53 7.39 8.82 1.46 Cactus Guard Hair 8.54 9.34 21 2.87 Underfur 5.74 7.97 17.84 2.32 Deer Guard Hair 6.23 7.75 8.95 3.06 Underfur 4.75 6.44 10.82 4.29 Pinyon Guard Hair 5.76 6.69 7.18 1.78 Western Harvest Guard Hair 9.12 13.39 15.73 4.06 Underfur 7.47 9.73 13.86 1.31 Northern Grasshopper Guard Hair 8.49 10.39 12.76 6.12 Southern Grasshopper Guard Hair 6.40 7.49 8.28 3.95 Dark Kangaroo Guard Hair 7.23 7.97 9.17 3.36 Little Pocket Guard Hair 6.86 8.38 9.68 8.14 Great Basin Pocket Guard Hair 8.36 12.94 16.75 2.39 Underfur 7.71 8.97 10.37 1.14 Silky Pocket Guard Hair 7.29 8.15 9.45 5.23 Long-tailed Pocket Guard Hair 5.86 6.83 7.82 5.59 Underfur 3.98 4.84 5.44 1.87 Desert Pocket Guard Hair 7.53 8.54 10.21 10.1 Rock Pocket Guard Hair 6.47 7.95 8.88 4.18 Western Jumping Guard Hair 6.00 7.78 9.41 2.79 Underfur 7.34 8.47 9.68 1.90 House Guard Hair 6.93 7.65 8.87 6.56 Underfur 10.23 12.83 15.58 2.51 Olive-backed Pocket Guard Hair 5.52 6.87 7.63 24.03 Underfur 12.37 14.40 20.74 6.06
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Figure 54: Boxplot of Guard Hair Widths (µm).
Figure 55: Boxplot of Guard hair Heights (µm).
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Figure 56: Boxplot of Underfur Widths (µm).
Figure 57: Boxplot of Underfur Heights (µm).
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CONCLUSION
Characteristic mammalian guard hair and underfur, as indicated by previous studies 23, Sessions et al 3, were found throughout the Myodoita and Castorimorphia suborder samples. The hair morphology characteristics of scale direction, scale pattern, hair width, and hair height were useful in characterizing mice between the families and even within the families. Unlike bat hair, an example of mammalian hair with immense variation within a single species but small variation within the family, mouse hair had small variation within species but greater variation between families and sub-orders 3. The most common scale pattern was the elongate-broad petal for the family Cricetidae; regular wave pattern for the family Heteromyidae; jagged elongate and joint-cusp for the family Muridae; and regular wave for the family Dipodidae. Greater accuracy in determining patterns for each family, however, could be achieved by increasing the sample size of the specimens, especially for the family Dipodidae where only one species was studied. It was observed that mice in the genus Peromyscus had a concavo-convex 26 or U-shaped, guard hair. It is possible that this feature could contribute to heat preservation or water repulsion. Due to the shaft shape, higher density and greater insulation could be achieved and maintained. In addition, maximizing the water contact surface area allows for optimal evaporation and run off. Cross-sectional hair morphology may be useful in elucidating further U-shape implications as earlier studies indicate 7, 8. Previous mouse hair studies used SEM and optical imaging of only a few species 28, 29. Our study, however, was large in comparison, including 18 species of indigenous Utah mice. Due to the greater sample size, we were more easily able to compare and contrast differences and similarities between families and suborders. Additional studies could further expand the sample size to include a more comprehensive compilation of other Rodentia species, including those outside of Utah. Tables 1-3 allow for correct identification of all mice species, which can be especially useful for forensics, archaeology, and taxonomy purposes (mammalian distributions).
ACKNOWLEDGMENTS
We thank Betsy Spackman for conducting scientific literature searches, Michael Standing for assistance with electron microscopy, and Duke Rogers for collecting the hair samples.
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Corresponding Author: Brad Carmack;
Department of Biology, Brigham Young University, Provo, UT, US
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