World Journal of Zoology 10 (2): 112-117, 2015 ISSN 1817-3098 IDOSI Publications, 2015 DOI: 10.5829/idosi.wjz.2015.10.2.93124 Tail Allometry of the Grasscutter (Thryonomys swinderianus) and African Giant Pouched Rat (Cricetomys gambianus): It s Functional Relevance 1 2 3 Byanet Obadiah, Tavershima Dzenda and Obadiah I. Happiness 1 Department of Veterinary Anatomy, College of Veterinary Medicine, University of Agriculture, Makurdi, Nigeria 2 Department of Veterinary Physiology, Faculty of Veterinary Medicine, Ahmadu Bello University, Zaria, Nigeria 3 Department of Biological Sciences, Faculty of Science, Benue State University, Benue, Nigeria Abstract: Tail dimensions and morphology are considered important traits for interpreting the functional adaptation of rodents to their ecological enclave. In this study, quantitative analyses of the body and tail dimensions were conducted in 20 African giantrats, AGRs (9 bucks and 11 does) and 12 greater cane rats or grasscutters, GCs (6 bucks and 6 does). The results showed that mean body length in males (bucks) was longer (P < 0.05) than that of females (does), but body circumference, tail length and weight differ not (P > 0.05) between sexes in both rodent species.with the exception of body circumference and width of tail tip, the mean body and tail lengths, tail weight, width of tail base and tail midpoint were significant between species (P < 0.05), with values higher in the AGR than the GC. The tail length and weight in the AGR, in particular, were almost three times (P < 0.0001) thevalues in the GC.In both rodents, the whole body length correlated positively (P < 0.01) with body and tail lengths.in the GC, the whole body length also correlated with tail weight (r = 0.8456, P < 0.0001) and width of tail tip (r = 0.6487, P < 0.01).In conclusion, bucks had longer body lengths than does, but similar tail lengths, in both species.longer and heavier tails, suggestive of more efficient thermoregulative and balancing abilities, were observed in the AGRs compared to the GCs. Key words: Tail Morphometric C. gambianus T. swinderianus INTRODUCTION to destruction of the ecosystem through the setting of bush fire by hunters [1]. The AGR climbs trees, using its The African giant rat (AGR), Cricetomys gambianus long, strong tail for balancing, whereas the GC is a poor and the greater cane rat or grasscutter (GC), Thryonomys climber with a short, fragile tail[1]. swinderianus are among the largest African fodents. Small nocturnal mammals, such as the AGR and Both rat species are nocturnal and found throughout GC, stay in burrows or holes during in day time to avoid tropical or sub-saharan Africa [1]. Animals in the exposure to heat during the hottest times of the day [8]. tropical forest have different adaptive measures to the The rodents possess some capacity to modify their environmental conditions as well as diets [2-4]. behavioural and internal body physiological features in The average weights in the GC were noted to be 9kg response to changes in environmental temperature [9]. for males (bucks) and 5 to 7 kg for females (does) [5], The rat tail, for example, has been shown to serve a aheatwhile that of the AGR is 1 to 1.6kg with body length of loss organ, providing a thermoregulatory function [10-11]. 400 mm [1, 6]. Both rodents inhabit the Sudan and Guinea Tail length was shown to be shorter in ground squirrels, Savannah agro-ecological zones, but the AGR is also longer in tree squirrels and longest in flying squirrels, found in the rain forest[7]. There is high demand for their having strong relationships with their ecological system meat and the economic benefit that accrues from their [12]. Sexual dimorphism in arboreal squirrels was also sales has resulted in aggressive hunting, for them leading observed, where females had longer tails relative to body Corresponding Author: Byanet Obadiah, Department of Veterinary Anatomy, College of Veterinary Medicine, University of Agriculture, Makurdi, Nigeria. 112
length than their male counterparts[12]. Thus, tail morphology is important for interpreting functional adaptation and phylogeny[13]. Tailless rats, a product of genetic mutation [14], experimentally showed limited capacity for thermoregulation compared to tailed rats [15]. Research on body-brain encephalization quotient indicated that the GC was more intelligent than the AGR [16], but the authors observed that the former was less able to adjust to rapid changes in environmental temperature. The GCs were usually transported very early in the morning or late in the evening, using cages constructed of light wood or plywood with several round holes for proper ventilation, in order to avoid heat stress and prevent high mortality. There is paucity of information on quantitative comparative analysis of the tail and body dimensions of AGRs and GCs in relation to their ecological adaptation. The goal of the present study was to quantitatively assess the patterns of body length and tail dimensions between AGRs and GCs and to investigate sexual dimorphism, with a view to relate these traits to their ecology and behaviour. MATERIALS AND METHODS Animal Source and Study Location: A total of 32 rodents, made up of 20 AGRs (9 bucks and 11 does) and 12 GCs (6 bucks and 6 does) were used in the study. The AGRs were live-trapped (using local traps) in the wild, area of Zaria and reared under laboratory conditions [13] in the Department of Veterinary Physiology and Pharmacology, Ahmadu Bello University, in the Northern Guinea Savannah zone of Nigeria. They were moved in laboratory cages to a nearby Veterinary Anatomy Research Laboratory, where the research was conducted. Matured GCs[18] were purchased from a breeder farm in Otukpo, Benue, Southern Guinea Savannah zone of Nigeria. They were then transported by road using wooden laboratory cages measured 50 cm (height) by 40cm (width) and 40cm (length), to the Anatomy Research Laboratory, Department of Veterinary Anatomy, Ahmadu Bello University, Zaria, Nigeria, where they were kept under room temperature for three days before the experiments began.the GCs were fed elephant grass (Pennisetum purpureum), supplemented with supplemented with chick grower mash and given access to water and feed ad libitum. The rats were anesthetised with 10-25 mg/kg IM ketamine HCl before euthanized. The body and tail dimension parameters were measured using twine, meter rule and digital vernier caliper (MG6001DC, General Tools and Instruments Company, New York). The parameters included; i) whole body length (WBL): distance from the tip of the nose to the tip of the tail, ii) body length (BL): from the tip of the nose to the caudal aspect of the body close to the base of the tail, iii) body circumference (BC): distance round the body at the flank area close to the rib cage, iv) tail weight (TW): weighed with a mettler balance (Model P 1210, AG, Switzerland, with sensitivity of 0.01g), v) tail length (TL): distance from base of tail to the tip, vi) width of the tail base (WTB), vi) width of tail midpoint (WTM) and vii) width of the tail tip (WTT), diameter were taken at the base, middle and tip of the tail, respectively. Data Analyses: Data collected were tabulated and expressed as mean and standard error of the mean (Mean ± SEM). All Statistical analyses were performed using the Statistical Analysis Package (Graph Pad Prism, Version 3.10, 2009. Student t-test was used to analyse the mean differences of tail and body dimensions between males and females of the same species and between species. Pearson's Correlation Coefficient (r) was used to compare the relationships between all the body and tail dimensions. Multiple regression tests were used to compare between the pairs of variables to determine their P values. Values of P < 0.05 were considered significant. RESULTS Table 1 displays the sexual dimorphism of the body and tail dimensions of AGRs and GCs.The mean WBL recorded in AGR bucks (807.8 ± 16.1 mm) was slightly (P = 0.0595) longert han that obtained in the does (764.5 ± 14.1 mm) and the parameter showed no sex difference (P = 0.5296) in the GCs. However, the BLs in both AGR and GC bucks (387.2 ± 8.4 mm and 318.2 ± 10.3 mm, respectively) were significantly (P < 0.05) longer than those of their female counterparts (352.4 ± 9.0 mm and 311.2 ± 32.2 mm, respectively). The rest of the measured body and tail dimensions (BC, TL, TW, WTB, WTM and WTT) did not differ (P > 0.05) between the sexes in both rodent species. Table 2 shows the comparison between the body and tail dimensions in the AGR and the GC. It was noted that out of the 8 parameters that were recorded, only the mean values of BC and WTT did not differ significantly (P > 0.05) between the 2 rodent species. The rest of the measured body and tail dimensions (WBL, BL, TL, TW, WTB and WTM) were significantly higher (P < at least 0.05) in AGRs than in GCs. In fact, the values of WBL, 113
Table 1: Sexual dimorphism in the body and tail dimensions of the African giant rat and grasscutter Male (n = 9) Female (n = 11) ---------------------------------------------- --------------------------------------------- Species Parameter Min-Max Mean ± SEM Min-Max Mean ± SEM P value African giant rat (n = 20) WBL (mm) 700 860 807.8 ± 16.1 650 810 764.5 ± 14.1 0.0595 BL (mm) 350 423 387.2 ± 8.4 305 405 352.4 ± 9.0 0.0125* BC (mm) 235 300 270.4 ± 7.3 265 320 280.7 ± 4.7 0.2591 TL (mm) 350 465 420.6 ± 10.6 320 450 412.1 ± 12.3 0.6090 TW (g) 450 600 522.2 ± 20.6 400 800 581.8 ± 46.4 0.2611 WTB (mm) 137.3 178.5 155.7 ± 4.0 139.1-200 158.3 ± 4.9 0.6908 WTM (mm) 71.0 96.8 87.2 ± 2.9 74.5 97.7 86.1 ± 2.3 0.7873 WTT (mm) 15 18.3 16.5 ± 0.4 14.3 22.1 17.9 ± 0.9 0.1693 Male (n = 6) Female (n = 6) Grasscutter (n = 12) WBL (mm) 404 535 451.5 ± 18.5 375 695 463 ± 47.7 0.5296 BL (mm) 294 375 318.2 ± 10.3 259 470 311.2 ± 32.2 0.0198* BC (mm) 150 365 242.5 ± 29.7 205 435 323.3 ± 40.9 0.1444 TL (mm) 110 160 133.3 ± 8.0 116 225 151.8 ± 15.5 0.3242 TW (g) 100 400 216.7 ± 60.1 50 600 187.5 ± 85.1 0.7864 WTB (mm) 452 161.8 100.5 ± 19 12 184.7 118.9 ± 24.2 0.5646 WTM (mm) 15.4 54.2 42.7 ± 9.2 4-76.6 46.3 ± 10 0.7970 WTT (mm) 17.6 22.9 20.9 ± 1.2 6 31.4 18.8 ± 3.3 0.5717 WBL: whole body length, BL: body length, BC: body circumference, TW: tail weight, TL: tail length, WTB: width of tail base, WTM: width of tail midpoint, WTT: width of tail tip, * = significant difference. Table 2: The body and tail dimensions in the African giant rat and grasscutter African giant rat (n = 20) Grasscutter (n = 12) -------------------------------------------------- ------------------------------------------------------ Parameter Min-Max Mean ± SEM Min-Max Mean ± SEM P value WBL (mm) 650 860 784 ± 11.4 375 695 457.3 ± 24.5 0.0001*** BL (mm) 330-395 368.1 ± 3.3 265 470 179.4 ± 15.7 0.0001*** BC (mm) 235-320 276.3 ± 4.0 150 435 277.9 ± 26.3 0.9531 TL (mm) 320 465 415.9 ± 8.1 110 225 142.6 ± 8.8 0.0001*** TW (g) 400 800 55.5 ± 2.7 50 600 20.2 ± 5.0 0.0001*** WTB (mm) 137.3 200 157.1 ± 3.2 120 184.7 113 ± 14.1 0.0101* WTM (mm) 71 97.7 86.6 ± 1.8 40 76.6 44.5 ± 5.6 0.0001*** WTT (mm) 14.3-22.1 17.3 ± 0.5 6 31.4 19. 6 ± 1.7 0.2187 WBL: whole body length, BL: body length, BC: body circumference, TW: tail weight, TL: tail length, WTB: width of tail base, WTM: width of tail midpoint, WTT: width of tail tip, * = significant difference, *** = very highly significant difference. BL, TL, TW and WTM were highly significantly (P < BL correlated with TW (r = 0.7970, P < 0.001); 0.0001) higher in AGRs. The values of TL and TW TL correlated (P < 0.05) with TW (r = 0.6264), WTM obtained in the AGR, for example, were almost thrice (r = 0.6780) and WTT (r = 0.6897); TW also correlated (P< 0.0001) those recorded in the GC. (P < 0.05) with WTB (r = 0.6684) and WTT (r = 0.6114); The results of correlation matrix computed for both WTB also correlated (P < 0.01) with WTM (r = 0.7747) and species (AGRs and GCs) are shown in Table 3. In both WTT (r = 0.7161); and WTM also correlated with WTT rodents, the WBL correlated positively (P <at least 0.01) (r = 0.8238, P < 0.01). All other correlations, including with BL and TL. In the GC, the WBL also correlated with those involving BC in both species were not significant TW (r = 0.8456, P < 0.0001) and WTT (r = 0.6487, P < 0.01); (P > 0.05). 114
Table 3: Correlation coefficients between the body and tail dimensions in the African giant rat and grasscutter Correlated Coefficient (Pearson r) ----------------------------------------------------------- Correlated Parameters African Giant Rat (n = 20) Grasscutter (n = 12) WBL and BL 0.7067*** 0.9487*** WBL and BC -0.1360 0.4248 WBL and TL 0.7746*** 0.7275** WBL and TW 0.2019 0.8456*** WBL and WTB 0.0614 0.5639 WBL and WTM 0.0269 0.5215 WBL and WTT -0.0525 0.6487* BL and BC -0.0624 0.4746 BL and TL 0.0999 0.4732 BL and TW 0.2124 0.7970** BL and WTB -0.2219 0.4588 BL and WTM 0.0465 0.3571 BL and WTT -0.0708 0.5150 BC and TL -0.1356 0.1537 BC and TW 0.1648 0.2693 BC and WTB 0.1518 0.0851 BC and WTM 0.0971 0.0991 BC and WTT 0.4913-0.1617 TL and TW 0.0942 0.6264* TL and WTB 0.1120 0.5755 TL and WTM -0.0037 0.6780* TL and WTT -0.0106 0.6897* TW and WTB -0.0042 0.6684* TW and WTM 0.1974 0.4434 TW and WTT 0.4344 0.6144* WTB and WTM 0.4534 0.7747** WTB and WTT 0.3510 0.7161** WTM and WTT 0.1181 0.8238** WBL: whole body length, BL: body length, BC: body circumference, TW: tail weight, TL: tail length, WTB: width of tail base, WTM: width of tail midpoint, WTT: width of tail tip, * = P < 0.05 = significant correlation, ** = P < 0.01 = very significant correlation, *** = P < 0.001 = very highly significant correlation. DISCUSSION The rodent tail as observed in AGRs and GCs in this work is an extension of the coccygeal vertebrae which projects at the caudal aspect of the animal. The tail is one of the distinguishing features of rats[19]. The tail is a long cylindrical structure, composed of three layers; the inner core made up of vertebrae, followed by a layer of tendons and a layer of skin externally. Blood vessels run the length of the tail between the tendons and aid in heat regulation[10]. Of all the body and tail dimensions measured in the present study, sexual dimorphism was demonstrated only with respect to the BL in both rodents. Bucks had significantly (P < 0.01) longer body lengths than their female counterparts in both species, although absolute TL did not differ between the sexes, indicating that does (females) had higher TL to BL ratios. The functional implication of the finding is that the does may possess better balancing and thermoregulatory abilities than bucks in both species[20]. The BL, TL and TW in the AGRs were significantly (P < 0.0001) longer or heavier than in the GCs. There was significantly positive correlations between the TL with TW (r = 0.0.6264, P < 0.01), WTT (r = 0. 6780, P < 0.01) and WTT (0.6897, P < 0.01) in the GC.Similar to our findings in GCs, the Norway rat TL (150 225mm) was shorter than BL (180 250mm [19]. Tail dimension may be a species variation or related to ecosystem. It was reported that TL in many species of squirrels was related to ecology; where the longest tail was observed in flying squirrels, followed by longer tails in tree squirrels and shorter tails in the ground squirrels [12]. Functionally, the tail aids in climbing and jumping abilities [14]. Therefore, the longer tail in AGRs may be an adaptive feature for better thermoregulatory, climbing and balancing abilities compared to the small and short tail of GC, with thermal intolerance and short range of climbing ability. The longer tail length observed in the AGR suggests[13] that the number of coccygeal vertebrae that make up the tail is higher in AGR than in GC, the later containing 18-23 bones [21]. To further look at tail function, [13] reported that tail in rat does not aid only in balancing, but also in counterbalances, enabling rats to stand upright with their hindlimbs and free their forelimbs in the air when feeding.the rats clamp ropes, walk along the fences and branches with tail performing the function of balancing and helps also to increase the rat rotational inertia, preventing rotation around the rope and can change its centre of gravity by the role of tail [10]. Based on the report of [22] and our personal experiences with these animals, the AGR has a very strong and long tail that is usually suspended in the air when running, unlike the GC with small, short and weak tail that can easily break in the process of handling. Thus, the AGR can climb trees using its long tail for balancing, unlike the short tail of GC, which may provide little or no appreciable support for balancing to the animal[1]. The findings agree with those of Larson and Stern [23], who reported that primate tail played an important role in maintaining balance in an arboreal environment. 115
The rat's tail has a thermoregulatory function, Anatomy, Faculty of Veterinary Medicine, Ahmadu Bello serving as a heat-loss organ. In hot environments, animals University, Zaria, Nigeria. This work was partly supported with long tailsmay be at an advantage, dissipating heat to by grants from Educational Trust Funds (ETF), Federal the environment as a consequence of the longer tail [24] Ministry of Education, Nigeria, through the University of better than animals with short tails. The tail performs this Agriculture, Makurdi, Benue State, Nigeria. function effectively due to its large surface to volume ratio and numerous blood vessels, especially at the middle REFERENCES and tip [25]. In the present study, the WTB and WTM were significantly wider in the AGR than in GC (P= 0.0101 1. National Research Council, USA, 1991. and P< 0.0001, respectively). This suggests higherblood Microlivestock: Little-Known Small Animals with a perfusion ability in the tail of AGRs than in GCs in Promising Economic Future. National Academy addition to its longer tail length; hence, better Press, Washington, D. C. pp. 225-240. thermoregulation in AGRs is expected.the finding may 2. Mona, N., J. Kamaruzama, R.B. Abdul and M. Nima, explain the observed high casualties recorded during road 2010. Modeling the potential distribution of wildlife transportation of GCs under hot conditions. Many GCs species in the tropics. World Journal of Zoology, have also been observed to die in the laboratory during 5(3): 225-231. the cold-dry (harmattan) season, suggesting that the GC 3. Susma, G.I, A. Achyut, K. Raj, B. Koirala and has poor thermoregulatory mechanism for dissipating or R. David, 2011. Feeding Ecology and Distribution of conserving heat under thermal stress. Himalayan Serow(Capricornisthar) in Annapurna The skeletal morphology of the coccygeal vertebrae, Conservation Area, Nepal.World Journal of Zoology, particularly the sacral and proximal caudal vertebrae are 6(1): 80-85. said to relate to tail length in most animals [13]. The 4. Ali, L.M., 2013. Macro-anatomical investigation of skeletal morphology of the GC has been documented [21], the skull of golden jackal (Canisaureus) and its but similar work in AGR is lacking in the available clinical application during regional anaesthesia. literature. Investigation also needs to be conducted on Global Veterinaria, 10(5): 547-550. the musculature of the tail, for its role in tail movements 5. Hoffmann, M., 2008. Thryonomys swinderianus. and strength, as well as blood vasculature. Also, a good The IUCN Red List of Threatened Species.Version technique for recording heat loss from the tail of these 2014.3.www.iucnredlist.org. Downloaded on 19 rodents in cold and heat periods may help to shed more February 2015 light on their thermoregulatory and climbing/balancing 6. Dzenda, T., J.O. Ayo, C.A.M. Lakpini and abilities. A.B. delaiye, 2011b. Seasonal and sex variations in live weights of captive African Giant rats CONCLUSION (Cricetomys gambianus, Waterhouse) in the Northern Guinea Savannah zone of Nigeria. It is concluded that sexual dimorphism was not International Journal of Zoological Research, observed in tail length, but males had longer body than 7(1): 49-58. their female s counterparts in both African giant pouched 7. Adenyo, C., A. Hayano, B.B. Kayang, E.H. Owusu rats and grasscutters. Comparatively, the longer, larger and M. Inoue-Murayama, 2013. Mitochondrial and heavier tails noted in the African giant pouched D-loop diversity of grasscutter rats is suggestive of better thermoregulatory and (Thryonom ysswinderianus Rodentia: balancing/climbing abilitiesthan grasscutters. The Hystricomorpha) in Ghana. Open Journal of Animal smaller size and shorter tails noted in the grasscutters Sciences, 3(3): 145-153. maybe associated with its observed relative inability to 8. Bennett, A., F. Raymond, B. Huey, J. Henry and cope with thermal stress in captivity. A.N. Kenneth, 2001. The parasol tail and thermoregulatory behaviour of the cape ground ACKNOWLEDGMENT squirrel xerusinauius. Physiology, 57(1): 57-62. 1984. 9. Rashed, R and M.A. El-Edel, 2015. Behavior and We acknowledge the technical assistance of the performance of growing rabbit under various floor staff of gross anatomy lab, Department of Veterinary types.global Veterinaria, 14(1): 149-155. 116
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