On Mammalian Sperm Dimensions J
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On mammalian sperm dimensions J. M. Cummins and P. F. Woodall Reproductive Biology Group, Department of Veterinary Anatomy, University of Queensland, St Lucia, Queensland4067, Australia Summary. Data on linear sperm dimensions in mammals are presented. There is infor- mation on a total of 284 species, representing 6\m=.\2%of all species; 17\m=.\2%of all genera and 49\m=.\2%of all families have some representation, with quantitative information missing only from the orders Dermoptera, Pholidota, Sirenia and Tubulidentata. In general, sperm size is inverse to body mass (except for the Chiroptera), so that the smallest known spermatozoa are amongst those of artiodactyls and the largest are amongst those of marsupials. Most variations are due to differences in the lengths of midpiece and principal piece, with head lengths relatively uniform throughout the mammals. Introduction There is increasing interest in comparative studies of gametes both from the phylogenetic viewpoint (Afzelius, 1983) and also in the analysis of the evolution of sexual reproduction and anisogamy (Bell, 1982; Parker, 1982). This work emerged as part of a review of the relationship between sperm size and body mass in mammals (Cummins, 1983), in which lack of space precluded the inclusion of raw data. In publishing this catalogue of sperm dimensions we wish to rectify this defect, and to provide a reference point for, and stimulus to, further quantitative work while obviating the need for laborious compilation of raw data. Some aspects of the material presented previously (Cummins, 1983) have been re-analysed in the light of new data. Materials and Methods This catalogue of sperm dimensions has been built up from cited measurements, from personal observations and from communication with other scientists. Where personal measurements are included, they were made on air-dried nigrosin-eosin live/dead sperm smears (Campbell, Dott & Glover, 1956) using a calibrated projection microscope. Some data are based on measurements made from Retzius' series of monographs (1906-1910). Although Retzius did not, unfortunately, give accurate scales for his illustrations, it appears that in most cases all the spermatozoa on a single plate were drawn to the same scale. On many plates, therefore, it was possible to find a reference spermatozoon of known dimensions, and to estimate the total sperm lengths of other species from this. While these measurements can be no more than rough approximations, they have nevertheless proved useful for comparative work. In Table 1, such measurements are only included when more accurate data for an otherwise unknown group are not available. When it subsequently proved possible to double-check such estimates, they were usually accurate to within 5 µ . The weights and classification of eutherian and monotreme mammals are from Walker (1975); marsupial taxonomy is from Kirsch (1977). Species identification is listed as given by the original source. Where necessary, more modern or more widespread synonyms are shown in parentheses. These were obtained from Ellerman (1940, 1941), Hall & Kelson (1959), Meester & Setzer (1974), Walker (1975) and Haltenorth & Differ (1980). Downloaded from Bioscientifica.com at 09/26/2021 06:19:00PM via free access Results and Discussion Table 1 gives a complete listing of all the data that we have collected on sperm dimensions. Except¬ ing only those figures keyed by an asterisk, which are probably inaccurate (see Key to Table 1), all figures have been accepted uncritically, and readers are referred to the original source for details of methodology, ranges and standard errors of mean measurements. Figures given in parentheses are estimates from Retzius' monographs (1906, 1909a, b, c, 1910: see 'Material and Methods'). Table 2 summarizes the extent of our data coverage. We now have collected quantitative data on 6-2% of all mammalian species, covering 17-2% of genera and 49-2% of families. We have been unable to find quantitative information for 4 minor orders; the Pholidota (pangolins), the Dermoptera (flying lemurs), the Sirenia (dugongs, manatees) and the Tubulidentata (aardvark). Of the larger orders, as might be expected, the extent of coverage is most comprehensive for those most likely to be used in the laboratory or as domestic species; the Artiodactyla, Perissodactyla, Lagomorpha and Primates. While we have data on more species of rodents (122) than for any other order, because of the enormous diversity and number of species, this represents only 5-3% of the total—slightly less than the average cover for all mammalian species. Marsupials and monotremes are reasonably well represented, reflecting much recent interest in the use of sperm morphology for phylogenetic analysis (see, for example Harding, Woolley, Shorey & Carrick, 1982). Rather surprisingly, there are no sperm dimensions published for American marsupials, even though details of morphology have been known since the beginning of the century. This account deals only with linear measure¬ ments, and not with other quantitative aspects such as mass, volume or density; for a detailed bibliography before 1964, together with an exhaustive discussion of sperm mensuration in terms of volume, mass and projected surface area, together with a discussion of the possible sources of error in measurement, see van Duijn (1975) and van Duijn & van Voorst (1971). For a review of the genetics of sperm dimensions in mammals see Beatty (1970, 1972, 1975), Williams, Beatty & Burgoyne (1970) and Burgoyne (1975). For correlating sperm dimensions with those of eggs, see Hartman (1929). Table 1. Linear dimensions (µ ) of mammalian spermatozoa Head Midpiece Principal piece Species (common name) Source Length Width Length Width length Total eutherian mammals Order Artiodactyla Family Bovidae Aepyceros melampus (impala) 1 6-52 4-13 7-63 3200 46-15 Antilope cervicapra (blackbuck) 2 48-4* Alcelaphus buselaphus (hartebeeste) I 4-82 3-06 7-23 36-32 48-37 Bos taurus 1 6-77 4-27 9-83 36-93 53-53 (ox) 3 50-80 4 2501* 61 8-79 4-62 -9-71 -4-72 70 10-2 5-4 Bubalus bubalis (water buffalo) 3 9-45 508 1419 33-35 56-99 Capra hircus (goat) 6 8-27 4-25 11-38 0-74 39-75 59-39 Capra ibex (ibex) 7 8-5 4-5 20-0 40-45 68-73-5 Downloaded from Bioscientifica.com at 09/26/2021 06:19:00PM via free access Table 1, cont. Head Midpiece Principal piece Species (common name) Source Length Width Length Width length Total Cephalophus abyssinicus ( = Sylvicapra grimmia abyssinicus) (common duiker) 5 [68] Connochaetes albojubatus ( = C. gnou) (black wildebeeste) [55] Connochaetes taurinus (blue wildebeeste) 501 3-56 7-86 33-90 46-77 Damaliseus dorcas (blesbok) 4-53 2-97 7-7 38-59 50-82 Gazella granii (Grant's gazelle) [48] Hemilragusjemlahicus (Himalayan tahr) 46-3* Litocranius walleri (gerenuk) [35] Ovis aries 5-78 10-27 45-69 (sheep) 8-2 140 40-45 62-2-67-2 Ovis musimon 311* (mouflon) 26-6* Ovis vignei (urial) 24-3* Tragelaphus strepsiceros (greater kudu) 6-41 3-75 8-99 30-14 45-54 Family Camelidae Camelas dromedarius (one-humped camel) 5-62 2-92 7-34 34-23 47-21 Family Cervidae Alces alces (moose) 10 8 5-2 54-7 60-65 Axis axis (chital) 2 40-6* Cervus elaphus (red deer) 5 [57] Odocoileus virginianus 11 10-0 60 70 350 52 (white-tailed deer) 8 18-9* Rangifer tarandus (reindeer) 12 7-636 5-027 10-38 38-48 56-496 Family Giraffidae Giraffa camelopardalis (giraffe) 13 5-77 3-77 47-77 Family Hippopotamidae Choeropsis liberiensis Opygmy hippopotamus) 2 31-5* Hippopotamus amphibius (hippopotamus) 1 3-77 2-65 7-85 21-87 33-49 Family Suidae Sus scrofa 14 8-5 4-2 100 361 54-6 (P'g) 1 6-55 3-47 11-88 21-53 39-96 71 84- 4-3- 9-4 5-5 Family Tayassuidae Tayassu tajacu [40] (collared peccary) 23-4* Downloaded from Bioscientifica.com at 09/26/2021 06:19:00PM via free access Table 1, cont. Head Midpiece Principal piece Species (common name) Source Length Width Length Width length Total Order Carnivora Family Canidae Canisfamiliaris 15 5-6 55-3 (dog) 4 14-54* 67 5-9 3-9 10-6 45-8 62-7 Vulpesfulva (American red fox) 16 8-0 5-0 600 Family Felidae Felis catus 4-76 2-56 8-34 0-77 46-25 59-35 (cat) 12-92* Panthera tigris ( = Leo tigris) (tiger) 17 5-80 3-59 43-63 Family Mustelidae Mêles meles (badger) 5 [80] Mustela vison (mink) 11 7-0 61 60 30-0 430 Family Ursidae Thalarctos maritimus (polar bear) 8 36-3* Ursus arctos (brown bear) 2 30-7* Family Viverridae Cryptoproctaferox (fossa) 5 [69] Eupleres goudotii (small-toothed mongoose) 5 [45] Paradoxurusjerdoni (Indian palm civet) 2 42-4* Family Procyonidae Ailuropoda melanoleuca (giant panda) 18 50 4-2 7-2 0-8 390 51-2 Order Cetacea Family Balaenopteridae Megaptera nodosa ( = M. novaeangliae) (humpback whale) 19 52-5-614 Family Delphinidae Globicephala melas (= G.melaena) Opilot whale) 5 [67] Phocoena communis ( = P. phocoena) (common porpoise) 20 73-8 Tursiops truncatus (Atlantic bottle-nosed dolphin) 21 4-5 2-0 4-0 560 64-5 Family Physeteridae Physeter catodon (sperm whale) 22 45-5 Downloaded from Bioscientifica.com at 09/26/2021 06:19:00PM via free access Table 1, cont. Head Midpiece Principal piece Species (common name) Source Length Width Length Width length Total Order Chiroptera Family Molossidae Mormopterus planiceps ( = Tadarida planiceps) (little mastiff-bat) 43 60 80 520 660 Family Phyllostomidae Anoura cultrata (long-nosed bat) 23 501 4-35 7-78 0-76 50-69 62-58 Artibeusjamaicensis (Jamaican fruit-eating bat) 23 3-80 341 8-28 0-62 Artibeus lituratus (big fruit-eating bat) 23 4-50 3-45 9-38 0-96 71-87 85-75 Carolila castanea (Allen's short-tailed bat) 23 4-79 3 95 9-9 0-88