鼩鼱科内6 个种的阴茎形态学

阴茎形态学特征是研究哺乳动物亲缘关系的有力工具,尤其是在近缘种之间。本试验选取鼩鼱科内的黑齿鼩鼱(Blarinella quadraticauda)、小纹背鼩鼱(Sorex bedfordiae)、大纹背鼩鼱(S. cylindricauda)、云南鼩鼱(S. excelsus)、四川短尾鼩(Anourosorex squamipes)和灰麝鼩(Crocidura attenuata)6 个种共31 号阴茎标本观察其阴茎形态学特征。研究结果表明:这6 个种的阴茎均缺乏阴茎骨;在6 个种的阴茎横切面上,尿道靠近阴茎腹侧面为阴茎海绵体包裹,没有单独的尿道海绵体存在,输精管由阴茎海绵体包裹,在阴茎末端形成两条输精管侧支;黑齿鼩鼱的阴茎具有独特的舌状结构,可作为鉴定黑齿鼩鼱的一个重要特征,鼩鼱属内的小纹背鼩鼱、大纹背鼩鼱和云南鼩鼱的阴茎头界限很难观察到,而黑齿鼩鼱、四川短尾鼩和灰麝鼩的阴茎头则具有明显的界限。     

Histomorphology of the glans penis in Vespertilionidae and Phyllostomidae species (Chiroptera,Mammalia)

The penises of bats are taxonomically distinctive in size and shape. In addition, they are variable in microscopic anatomy, indicating that histomorphological studies of copulatory organs of bats may help understanding their successful reproductive strategies. We studied adult males of 13 species of vespertilionid and phyllostomid bats. Both families exhibited the basic structure of the vascular penis of mammals: the hydrostatic elements of the corpora cavernosa and the corpus spongiosum surrounding the urethra, as well as accessory cavernous tissue. Variation in the position and amount of the tissues were observed in these families. Vespertilionid bats have a small glans penis with abundant accessory cavernous tissue on the prepuce and a highly variable baculum. The baculum varied in size and morphology, even among congeneric species, such as the three Lasiurus species and the two Myotis species. Phyllostomid species possess no bacula, but vascular structures are present to produce penile stiffening, particularly on the glans. Variation in the microscopic anatomy of the phyllostomid prepuce was observed, for example, Artibeus species had accessory cavernous tissue surrounded by a tunica albuginea, but Carollia perspicillata had two bundles of striated musculature and some adipose tissue; abundant pigments were present in the prepuce of most species.     

Anatomy of the baculum-corpus cavernosum interface in the norway rat (Rattus norvegicus), and implications for force transfer during copulation

The baculum is a nonappendicular bone found in the glans tissue of members of five orders of mammals. Its function during copulation is unknown. Anatomical examination of the baculum and corpus cavernosum in the Norway rat (Rattus norvegicus) shows that the two structures are connected by a layer of fibrocartilage, and that the distal tip of the corpus cavernosum swells during erection to surround the proximal end of the baculum. Microradiographs of bacula from sexually experienced males show that regions of the bone may be remodeling; these data suggest that the baculum is load-bearing. On the basis of this anatomy, I propose that the baculum increases the overall flexural stiffness of the penis during copulation by transferring bending and compressive forces from the distal end of the glans to the tensile wall of the corpus cavernosum. Forces on the distal end of the penis during copulation press the baculum against the corpus cavernosum, reducing its internal volume and increasing intracavernosal pressure and corpus cavernosum wall strains. Because the wall of the erect corpus cavernosum is reinforced with inextensible collagen fibers, an increase in wall strain will also increase wall tissue stiffness, and thereby increase the flexural stiffness of the corpus cavernosum.     

中国13种鼠科啮齿动物的阴茎形态学及其分类学意义的探讨

本文对我国所产13种鼠科啮齿类进行了阴茎形态学的比较研究(包括阴茎骨和软体结构),探讨了在鼠科的家鼠属(Rattus)、姬鼠属(Apodemus)、小家鼠属(Mus)和巢鼠属(Micromys)4个属间以及同属不同种之间的差异。结果表明,阴茎的形态结构具有属和种的特异性和稳定性,可以为分类学的研究提供重要依据。     

Penial differentiation in speciation of subterranean mole rats Spalax ehrenbergi in Israel

Geographic and ontogenetic variation was examined in the glans penes and bacula of 92 specimens representing four biological species with the karyotypes 2n=52, 54, 58 and 60 of the Spalax ehrenbergi superspecies in 13 populations from Israel and one disjunct population from northern Egypt. Five measured and three calculated penial characters were analysed and employed in multivariate statistics. Comparison was made between penial morphological characters and craniometric biochemical and chromosomal data. The baculum is strongly species-specific and its length bears no significant relationships with head and body length. The results of these analyses suggest that different bacular morphologies may provide a basis for species-specific prezygotic isolating mechanisms, thus, together with other factors, contributing to reproductive isolation and speciation.     

Baculum length and copulatory behavior in primates

The length of the baculum (os penis) was measured in 74 adult males representing 46 primate species. These data, and a review of previously published measurements, indicate that variation in baculum length among primates is related to taxonomic and behavioral differences. Thus, many New World monkeys have shorter bacula, relative to body weight, than Old World monkeys. The baculum is shorter in colobine monkeys than in cercopithecines. Among the great apes, reduction of the baculum is more pronounced in Pan and Gorilla than in Pongo. Very long bacula are found in some nocturnal prosimians (eg, Lorisidae) and also in Macaca arctoides. A comparison of baculum length relative to body weight was made in 34 species for which detailed information on copulatory behavi or was available. The presence of an elongated baculum was shown to correlate with copulatory patterns involving prolonged intromission and/or the maintenance of intromission during the postejaculatory interval (eg, Galago crassicaudatus, Loris tardigradus, M, arctoides). The evolutionary significance of these observations is discussed and it is suggeted that similar copulatory patterns may occur in species with elongated bacula (eg, Daubentonia, Perodicticus) for which behavioral data are lacking at present. The same hypothesis also applies to an extinct adapid primate which possessed a very large baculum.     

Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)

Relationships between baculum length, body weight and copulatory behaviour were examined in 66 species of carnivores and pinnipeds (Grand Order Ferae). Elongated bacula occur in most carnivore species of the families Ursidae, Canidae, Procyonidae and Mustelidae as well as in all pinnipeds studied. By contrast, members of the family Felidae have short bacula in relation to their body weights. Elongate bacula are found in carnivores and pinnipeds with a prolonged single intromission (PI) copulatory pattern. This finding agrees with results of a previous study of baculum length and PI copulatory patterns in primates. The enlarged baculum may serve to strengthen the penis and protect the urethra during prolonged intromissions. The distal pole of the baculum may also assist sperm transport since in some species it projects beyond the tip of the penis and probably contacts the female's os cervix during copulation. It is possible that stimulation of the female's genitalia by the baculum might also be important in mammals which are induced ovulators (e.g. Mustelidae). However, it is notable that elongation of the baculum has also occurred in some groups where females ovulate spontaneously (e.g. Canidae, Primates).     

沟牙田鼠的形态特征及分类地位探讨

沟牙田鼠是分布于中国四川西部和甘肃南部的一种罕见小型啮齿动物,迄今标本收藏甚少,种、属地位存在异议,有人认为应并入田鼠属。我们根据四川九寨沟自然保护区采到的16号不同年龄、性别的沟牙田鼠,就其牙齿形态和成年雄性的阴茎结构与其它田鼠类作了比较,发现其成体和幼体在M³和M₃形态上有很大不同。其他重要区别包括：上门齿宽大;下门齿外露部分很短,下门齿总长仅及下颌长的77％左右,远比其他田鼠小;M齿环呈圆弧形或豆形亦很特殊;成年雄性的阴茎骨近支烧瓶状,远支基部膨大与田鼠类差别很大。结果表明：沟牙田鼠属是有别于田鼠属的有效属。     

Male external genitalia and systematic relationships of the Japanese macaque (Macaca fuscata Blyth, 1875)

Penial morphology was studied in six preserved specimens of the Japanese macaque,Macaca fuscata. Structure of the glans penis and baculum inM. fuscata is essentially similar to that in the rhesus macaque,M. mulatta. With age the skin of the penis inM. fuscata apparently becomes progressively more darkly and extensively pigmented, and the baculum becomes progressively longer. One immature specimen examined, with penial skin unpigmented and baculum cartilaginous, appears to be abnormally retarded in penial development. Penial morphology suggests that the allopatric speciesM. fuscata, M. mulatta, M. cyclopis, andM. fascicularis constitute a closely interrelated group. Contrary to a previous report, there is no evidence that penial morphology inM. fuscata is transitional to that inM. radiata andM. sinica. InM. fuscata, M. mulatta, andM. fascicularis, latitudinal variation of body size and relative tail length conforms toBergmann's rule andAllen's rule; external proportions in three specimens ofM. cyclopis only partly conform to this pattern.     

长爪沙鼠阴茎头的组织学研究及五种鼠阴茎头的扫描电镜观察

本文对８例长爪沙鼠（Ｍｅｒｉｏｎｅｓｕｎｇｕｉｃｕｌａｔｕｓ）幼年、亚成年、成年和老年雄性个体的阴茎头组织进行连续切片。结果表明,阴茎头的外环层和阴茎骨近支由阴茎海绵体衍生而来;内环层、尿道小瓣、阴茎骨远支和侧支由尿道海绵体衍生而成。采用扫描电镜观察了长爪沙鼠、子午沙鼠（Ｍ．ｍｅｒｉｄｉａｎｕｓ）、大沙鼠（Ｒｈｏｍｂｏｍｙｓｏｐｉｍｕｓ）和甘肃绒鼠（Ｅｏｔｈｅｎｏｍｙｓｅｖａ）、黑腹绒鼠（Ｅ．ｍｅｌａｎｏｇａｓｔｅｒ）共５种１６例的阴茎头表皮棘,发现其形态、数量和分布有属、种间差异性和相对稳定性,沙鼠属（Ｍｅｒｉｏｎｅｓ）３种的表皮棘均呈牛角状的圆锥体,斜向排列成行;而绒鼠属（Ｅｏｔｈｅｎｏｍｙｓ）内甘肃绒鼠为球状表皮棘,黑腹绒鼠为牛角状,排列均不规则。因此,阴茎头表皮棘与阴茎骨、阴茎头软体结构均有分类学意义。     

The canine baculum: the structure and mechanical properties of an unusual bone

The baculum is an extraskeletal bone located in the penis of a few species in several orders of mammals such as carnivores, insectivores, rodents, bats and primates. This study aims to describe the structure, architecture and mechanical properties of the canine baculum. To this end canine bacula from castrated and uncastrated dogs were collected and examined by light microscopy, micro-computed tomography (microCT) scanning, histological staining, and mechanical testing. Their mineral density and mechanical properties were compared with those of a typical skeletal bone (the radius) in the same dog. Furthermore, a numerical model of a representative baculum was created and its mechanical performance analyzed using the finite element method, in order to try to elucidate its function. Examination of light microscopy images of transverse sections shows that the baculum consists of a typical sandwich structure, with two cortical plates separated, and joined, by loose cancellous bone. MicroCT scans reveal that the mineral density is lower in the baculum than in the radius, both in castrated as well as in uncastrated dogs, resulting in much lower stiffness. Castration was found to decrease the mineral density in both the baculum and the radius. The most likely function of the baculum of the dog is to stiffen the penis to assist intromission, and its much lower mineral density compared to that of the radius may be a mechanism designed to decrease the stiffness somewhat, and thus reduce the risk of fracture during copulation.     

The vasculature of the rat penis: a scanning electron microscopic and histologic study.

As a basis for understanding the mechanism of erection in an animal model frequently used in research in reproductive biology, the angioarchitecture of the penis of the rat has been described using scanning electron microscopy. Study of the penile vasculature of the rat indicates that the corpora cavernosa penis and the corpus spongiosum are independent erectile tissues, each with its own arterial and venous vessels. The large vascular spaces and abundant smooth muscle of the penile crura are compatible with its role in regulating blood flow to more distal penile tissues. Helicine arteries of the crura, but not the parent deep penile artery or arteries elsewhere, have muscular cushions in their walls. The venous drainage of the penile crura is via subtunical veins which are thought to be compressed during erection to elevate pressure within the penis. Large, paired cavernous veins drain the shaft of the penis. A unique method for inhibiting blood flow from the penis is indicated by the division of the cavernous veins into smaller channels prior to joining the subtunical venous plexus. Erectile tissue in the bifid origins of the corpus spongiosum has abundant cavernous muscle, while in the remainder of the corpus spongiosum little smooth muscle lines the cavernous spaces. The cavernous spaces on either side of the urethra coalesce to form vessels, each of which communicates with cavernous spaces in the glans. In addition, a bypass of the glans is effected by communication of these vessels directly with the deep dorsal vein. The apparent absence of muscular pads in vessels of the spongiosum, the relative paucity of cavernous smooth muscle, and the ample venous drainage provided by the deep dorsal vein may account for the lack of a venous occlusive mechanism similar to that of the corpora cavernosa penis.     

Allometry of the baculum and sexual size dimorphism in American martens and fishers (Mammalia: Mustelidae)

Genitalia are among the most variable of morphological traits, and recent research suggests that this variability may be the result of sexual selection. For example, large bacula may undergo post‐copulatory selection by females as a signal of male size and age. This should lead to positive allometry in baculum size. In addition to hyperallometry, sexually selected traits that undergo strong directional selection should exhibit high phenotypic variation. Nonetheless, in species in which pre‐copulatory selection predominates over post‐copulatory selection (such as those with male‐biased sexual size dimorphism), baculum allometry may be isometric or exhibit negative allometry. We tested this hypothesis using data collected from two highly dimorphic species of the Mustelidae, the American marten (Martes americana) and the fisher (Martes pennanti). Allometric relationships were weak, with only 4.5–10.1% of the variation in baculum length explained by body length. Because of this weak relationship, there was a large discrepancy in slope estimates derived from ordinary least squares and reduced major axis regression models. We conclude that stabilizing selection rather than sexual selection is the evolutionary force shaping variation in baculum length because allometric slopes were less than one (using the ordinary least squares regression model), a very low proportion of variance in baculum length was explained by body length, and there was low phenotypic variability in baculum length relative to other traits. We hypothesize that this pattern occurs because post‐copulatory selection plays a smaller role than pre‐copulatory selection (manifested as male‐biased sexual size dimorphism). We suggest a broader analysis of baculum allometry and sexual size dimorphism in the Mustelidae, and other taxonomic groups, coupled with a comparative analysis and with phylogenetic contrasts to test our hypothesis. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 955–963.     

EXPERIMENTAL EVIDENCE FOR THE EVOLUTION OF THE MAMMALIAN BACULUM BY SEXUAL SELECTION

Male genitalia exhibit a taxonomically widespread pattern of rapid and divergent evolution. Sexual selection is generally believed to be responsible for these patterns of evolutionary divergence, although empirical support for the sexual selection hypothesis comes mainly from studies of insects. Here we show that sexual selection is responsible for an evolutionary divergence in baculum morphology among populations of house mice Mus domesticus. We sourced mice from three isolated populations known to be subject to differing strengths of postcopulatory sexual selection and bred them under common‐garden conditions. Mice from populations with strong postcopulatory sexual selection had bacula that were relatively thicker compared with mice from populations with weak selection. We used experimental evolution to determine whether these patterns of divergence could be ascribed to postcopulatory sexual selection. After 27 generations of experimental evolution, populations of mice subjected to postcopulatory sexual selection evolved bacula that were relatively thicker than populations subjected to enforced monogamy. Our data thereby provide evidence that postcopulatory sexual selection underlies an evolutionary divergence in the mammalian baculum and supports the hypothesis that sexual selection plays a general role in the evolution of male genital morphology across evolutionary diverse taxonomic groups.     

High diversity of pipistrelloid bats (Vespertilionidae: Hypsugo,Neoromicia, and Pipistrellus) in a West African rainforest with the description of a new species

The pipistrelloid bats (genera Hypsugo, Neoromicia, and Pipistrellus) of Africa have been poorly studied, partly as a result of problems associated with species identification. This paper examines the diversity of pipistrelloid bats from Mount Nimba, a biodiversity hotspot in the Upper Guinean rainforest zone. Traditional morphometrics, the structure of the baculum, and sequences of the cytochrome oxidase subunit I (COI) gene were used to identify taxa. Species richness was exceptionally high and included at least ten taxa identifiable on molecular grounds. Of these, existing names could be assigned to six taxa. A seventh taxon was described as a species new to science, Neoromicia roseveari sp. nov. , and was distinguished on molecular grounds, craniodental morphology, and baculum structure. The remaining taxa may refer to as‐yet undescribed species but we lacked sufficient material to formally describe them here. The high species richness of pipistrelloid bats on Mount Nimba may be associated with the transition zone from lowland rainforest to moist savannah. © 2013 The Linnean Society of London     

Are long penis bones an adaption to high latitude snowy environments?

Animals adapted to high latitude environments typically live at relatively low densities and require large areas to acquire the requisites for successful reproduction. Also, the unpredictable distribution of energy across time and space, such as environmental seasonality, constrains the timing of mating and parturition and imposes limitations to reproductive strategies. For example, limited mobility due to snow accumulation may reduce mate encounters, and this selective pressure may have resulted in morphological adaptations to assess mate quality during serial encounters. We tested whether mammalian carnivores living in high latitude environments displayed greater use of copulatory competition via length of the penis bone or baculum, and whether females in such settings enhance competition among males via multi-male matings. We then examined how baculum length may correlate with other reproductive adaptations in high latitude environments such as delayed implantation and induced ovulation. Statistical methods that account for phylogeny revealed that longer bacula occur in carnivores living in high latitude environments with greater snow accumulation, and that larger bacula is also associated with delayed implantation and multi-male mating systems. In contrast, the smallest bacula were observed in polygynous species. Our results provide an example of the effects of ecology on the evolution of a secondary sexual character due to sexual selection. Selection pressures imposed by seasonality of resources and environmental conditions affecting locomotion may have led to sexual selection via mating systems for females, and counter morphological adaptations of long penis bones in males.     

Penile Morphology and Classification of Bush Babies (Subfamily Galagoninae)

The penile morphologies of nocturnal prosimians are complex and vary considerably between genera and species. Accordingly, comparative morphology can be useful in taxonomic studies, particularly when assessing the status of newly discovered species. I measured features of penile morphology—surface area of the glans penis; shape and size of the keratinized spines on the glans—for populations representing 14 species within the subfamily Galagoninae. Intraspecific variations in penile morphology were relatively minor. By contrast, there are significant differences in several morphological features among closely related, sympatric species, e.g., in the greater bush babies (Otolemur crassicaudatus and O. garnettii) and lesser bush babies (Galago senegalensis and Galago moholi). Assessment of glans area resulted in the recognition of a second needle-clawed form: Euoticus pallidus. Similar divisions exist in the dwarf and greater bush babies with respect to proportional spiny area and characteristics of spine size. I constructed a key based on the presence/absence of certain features—penile spines, dermal markings on the glans, penile lappets—as well as the shape of the baculum and possession of different spinal morphotypes. This key may be used to identify all 14 species of bush babies. Penile morphologies provide a useful guide to specific identity in the Galagoninae, which may be true also for other groups of nocturnal mammals.     

The baubellum is more developmentally and evolutionarily labile than the baculum

Understanding the evolutionary forces that influence sexual dimorphism is a fundamental goal in biology. Here, we focus on one particularly extreme example of sexual dimorphism. Many mammal species possess a bone in their penis called a baculum. The female equivalent of this bone is called the baubellum and occurs in the clitoris, which is developmentally homologous to the male penis. To understand the potential linkage between these two structures, we scored baculum/baubellum presence/absence across 163 species and analyzed their distribution in a phylogenetic framework. The majority of species (N = 134) shared the same state in males and females (both baculum and baubellum present or absent). However, the baubellum has experienced significantly more transitions, and more recent transitions, so that the remaining 29 species have a baculum but not a well‐developed baubellum. Even in species where both bones are present, the baubellum shows more ontogenetic variability and harbors more morphological variation than the baculum. Our study demonstrates that the baculum and baubellum are generally correlated across mammals, but that the baubellum is more evolutionarily and developmentally labile than the baculum. The accumulation of more evolutionary transitions, especially losses in the baubellum, as well as noisier developmental patterns, suggests that the baubellum may be nonfunctional, and lost over time.     

Constructed wetlands support bats in agricultural landscapes

Bats are known to use aquatic habitats as foraging habitats. Agricultural intensification is perceived to be a main reason for the loss of wetlands. However, artificial wetland creation (i.e. the construction of retention-ponds) in the agricultural landscape aiming at water or nutrient retention has recently gained importance. We evaluated to what extent bats use these artificial wetlands as foraging habitats in an agricultural landscape.Bat activity and prey density were compared in matched pairs at retention-ponds and neighbouring vineyard sites using stationary bat-detectors and sticky-traps, respectively. To examine if bat activity is related to the number of bat individuals, a thermal infrared imaging camera was used. Pipistrellus pipistrellus, the dominant species, served as an example to assess habitat selection between retention-ponds and vineyards. This was performed by relating foraging activity to the available area available within the potential home-range.Total bat activity and nocturnal prey density were significantly higher above the retention-ponds than above vineyards. High differences of activity levels between the ponds and the respective vineyard sites were found for Pipistrellus spp. (P. pipistrellus and P. nathusii) and Myotis spp. (M. daubentonii and M. mystacinus), being about 180 times and 50 times higher above the retention-ponds, respectively. A significant correlation was found between recorded bat activity and the maximum number of bat individuals observed with a thermal infrared imaging camera. When relating foraging activity to habitat availability within the assumed home-range of P. pipistrellus, retention-ponds had on average a higher importance as a foraging habitat than the complete vineyard area although they covered less than 0.1% of its area.This study indicates that artificial wetlands such as retention-ponds provide foraging habitats for bats. Therefore, creation of wetlands in intensively used agricultural landscapes benefits bats.     

柔裳蜉属亚洲种类比较及紫金柔裳蜉稚虫再描记（蜉蝣目：细裳蜉科）

柔裳蜉属Habrophlebiodes蜉蝣的亚洲种类因两阳茎在端部明显分叉而与北美种类有显著区别。本属在亚洲地区现已知4种,其中出众柔裳蜉H.prominens Ulmer的下唇须末节较长、后翅脉相独特以及翅面着色明显而易识别;吉氏柔裳蜉H.gilliesi Peters阳茎较细,两阳茎叶的基部2/3愈合;紫金柔裳蜉H.zijinensis You&Gui阳茎短粗,两阳茎叶在基部1/2愈合,端部突起也较大。与吉氏柔裳蜉相比,紫金柔裳蜉稚虫的鳃具有多而明显的气管分支;与娇弱柔裳蜉H.tenella Kang&Yang相比,紫金柔裳蜉的鳃分叉较深、气管分支更近基部,体色也较浅。柔裳蜉属稚虫的主要特征与拟细裳蜉属Paraleptophlebia和新细裳蜉属Neoleptophlebia相似,但其中舌叶具细毛簇。本属所在的细裳蜉亚科Leptophlebiinae各属之间的系统发育关系重建需要对更多材料进行更深入的研究。