Lactation length in three species of Galago

Milk expression was used to determine lactation length in three species of Galago at the Duke University Primate Center. Lactation length in G. senegalensis moholi was significantly shorter than the lactation length of either G. garnettii or G. crassicaudatus, but lactation length did not differ between G. garnettii or G. crassicaudatus. Litter size had no significant effect on lactation length. The lactation lengths determined in this study are substantially longer than those reported by investigators who used observational techniques. These discrepancies may be partially explained by failure to observe nursing during the diurnal inactive period.     

Perinatal size and maturation of the olfactory and vomeronasal neuroepithelia in lorisoids and lemuroids

Explanations for the chemosensory abilities of newborn mammals focus primarily on food (milk) acquisition and communication (e.g., maternal-infant bonding). However, the relative importance of the main and accessory (vomeronasal) olfactory systems is hypothesized to differ at birth between altricial and precocial mammals. Strepsirrhines (lemurs and lorises) possess main and accessory olfactory systems, and vary in life-history traits related to infant dependency and maturation. Accordingly, this study examines the size and maturational characteristics of vomeronasal (VNNE) and olfactory (OE) neuroepithelia in strepsirrhines. Serially sectioned heads of 18 infant cadavers were examined microscopically for neuroepithelial distribution. Measurements were taken on the length of the nasal fossa on one side that was occupied by VNNE and OE. The data were corrected for body size using the cranial length or body mass, and were then examined for correlation with several life-history variables, as well as activity pattern. In addition, immunohistochemistry was used to identify cells in the VNNE and OE that express olfactory marker protein (OMP), a marker of mature olfactory neurons. Relative OE extent was not significantly correlated with any of the life-history variables. Relative VNNE length was negatively correlated with relative gestation length and relative neonatal mass (P<0.05). However, when we corrected for phylogenetic relationships, we found no significant correlations between either of the neuroepithelial measurements and life-history variables. Immunohistochemical findings suggest that OE has more OMP-reactive cells than VNNE in all species. OMP-reactive cells appear to be less numerous in diurnal species compared to most nocturnal species. These results indicate that the VNNE may be relatively longer at birth in altricial species. However, it remains uncertain how phylogeny and/or ontogeny may explain these findings.     

Loud calls ofGalago crassicaudatus andG. garnettii and their relation to habitat structure

Long distance vocalizations have been shown to be good indicators of genetic species in primates. Here the loud calls of two recently identified greater galago taxa —Galago crassicaudatus andG. garnettii — are compared and analyzed statistically. Observed differences in call structures are investigated further as potential indicators of differences in the structures of habitats frequented by the two species. Although the calls share a repetitive structure, and show similar dominant frequency bands (1,000 – 1,500 Hz), they differ significantly in the number of units per call, unit duration, inter-unit interval, highest frequency, lowest frequency, dominant frequency band, first harmonic, and call duration. The duration of theG. crassicaudatus call is more than twice that ofG. garnettii. Strong intraspecific consistency is seen in the most energetic frequency bands (dominant frequency band and first harmonic), and durations of the individual units and inter-unit intervals. Information important to species recognition is thus most likely to be contained in these features. Individual recognition may be encoded in the relative emphasis of higher level harmonics. The frequency structures of the calls will reflect requirements for acoustical transmission in a forest environment, as well as structural constraints imposed by body size. Higher frequencies detected in theG. garnettii call (up to 8,500 Hz) may have a functional significance related to distance estimation, or may simply be a reflection of smaller body size. The greater modulation of theG. garnettii call suggests that its habitat constitutes a denser or more turbulent medium for sound transmission than does the habitat ofG. crassicaudatus.     

Distribution of erythrocytic allozymes in two sibling species of greater galago [Galago crassicaudatus E. Geoffroy 1812 and G. garnettii (Ogilby 1838)]

This study investigated the use of erythrocyte enzymes as indicators of the presence or absence of gene flow between the sibling species G. crassicaudatus and G. garnettii. Fifty-five animals deriving from 14 different source populations were included in the analyses. In addition to hemoglobin, eight enzyme systems were examined: acid phosphatase, adenylate kinase, carbonic anhydrase II, esterase D, glucose-6-phosphate dehydrogenase, 6-phosphogluconate dehydrogenase, peptidase A, and peptidase B. of these, adenylate kinase, glucose-6-phosphate dehydrogenase, hemoglobin, peptidase A, and peptidase B showed no interspecific or intraspecific variation. Esterase D was polymorphic in certain populations of G. crassicaudatus but not in others or in G. garnettii. Acid phosphatase and 6-phosphogluconate dehydrogenase were polymorphic in G. garnettii but monomorphic in all G. crassicaudatus populations. The taxa showed fixation for different alleles at the carbonic anhydrase II locus, indicating a lack of gene exchange between the taxa. We suggest that acid phosphatase, 6-phosphogluconate dehydrogenase, and carbonic anhydrase II may be used as genetic markers in the identification of these two taxa.     

The use of hair morphology in the classification of galagos (Primates, subfamily galagoninae)

The outer epidermal hair structure of bushbabies or galagos varies considerably between genera and species. Comparative morphology can therefore be useful in taxonomic studies, particularly when assessing the true status of newly nominated or misclassified species. Quantitative measurements of outer epidermal hair scales (scale frequency and scale complexity) were made for populations representing 13 taxa within the subfamily Galagoninae. Intraspecific variations in both measures of hair morphology were relatively minor. By contrast, significant interspecific differences were observed, even between closely related, well-established species (e.g. in the greater galagosOtolemur crassicaudatus andO. garnettii and lesser galagosGalago senegalensis andGalago moholi). Assessment of scale frequency resulted in the recognition of all species of the greater and needle-clawed galago genera and added support for nomination to species status of two former subspecies (O. argentatus andE. pallidus). Similar divisions were made in the lesser and dwarf galagos with respect to scale complexity. Possible explanations for the differences in outer epidermal hair scales are discussed. Characteristics of outer epidermal hair scales provide a useful guide to species identity in the subfamily Galagoninae, and the same may prove to be true for other groups of nocturnal mammals.     

The use of hand morphology in the taxonomy of galagos

Measurements of volar (hand) pad area were made for 244 specimens, representing 12 species and 4 genera of galagos (sub-family Galaginae). When corrected for body weight, statistically significant differences were identified, at both the genus and species levels in the areas of the volar pads. Most informative, in terms of taxonomic differences, were measurements of two of the five pads; interdigjtal pad number 4 and thenar pad number 5. Closely related species were distinguishable on the basis of these measurements. The thick-tailed galago (Otolemur crassicaudatus) was separate from Garnett's galago (O. garnettii) and the specific status of the silver greater galago (O. argentatus) was supported. Likewise, the two needle-clawed galagos (Euoticus elegantulus andE. pallidus), recently separated on mitochodrial DNA grounds, were found to differ significantly in their volar pad morphology. These studies show that comparative studies of volar hand pad morphology provide a novel approach to the re-assessment of galago taxonomy, and may be applicable also in taxonomic studies of other prosimian groups.     

Speciation in the greater galagos (Prosimii: Galaginae): review and synthesis*

A marked morphological and genetical discontinuity among the greater galagos points to the existence of two genetical species within the taxon, Galago crassicaudatus E. Geofl'roy 1812. This contribution investigates the conditions surrounding this speciation event, and places it within a phylogenetic context. Information pertaining to body size, litter size, sexual dimorphism, locomotor mode and karyotypic structure is assessed. It is hypothesized that G. crassicaudatus diverged from its more gracile sibling, G. garnettii, approximately 2 million years ago, in respotise lo the increasing aridification of Africa that accompanied the Northern Hemisphere Pleistocene glaciations.     

Social organization of Galagos in Kenyan coastal forests: II. Galago garnettii

The social organization of Galago garnettii was studied for the first time and the study included data from two different sites in the coastal forests of Kenya. A combination of mark-recapture and radio-tracking techniques was used to investigate patterns of inter- and intrasexual home range overlap. Patterns of range use were established by radio tracking focal individuals. Adult females of different ages shared highly overlapping ranges, while like-aged females showed little range overlap. Females matured and had their first infants in their natal ranges. Adult males' ranges were larger than those of females and overlapped them extensively. Resident adult males showed little range overlap with each other, unless they were of different ages. Turnover of males was frequent in both populations. Males probably dispersed from their natal ranges. Adults rarely slept together. The social organization of G. garnettii is relatively similar to that of the closely related G. crassicaudatus of southern Africa but contrasts with that of its sympatric congener, G. zanzibaricus. The differences and similarities between these three species are discussed in relation to diet and body size.     

Palatal growth inMacaca nemestrina

A longitudinal study of the palatal growth of 80 male and 81 femaleMacaca nemestrina reveals that male growth rates are greater than female growth rates. In addition, both sexes demonstrate a similar pattern of palatal growth although growth rate differences between sexes probably reflect the larger canine/P₃ apparatus and overall body size of males. Growth of the premaxillary and postmaxillary palatal segments appear to be independent of one another. Palatal growth spurts, obvious in males, exist for two palatal length dimensions. The maxillary tuberosity growth center is postulated to account for more size dimorphism between sexes than the premaxilla growth center.     

Isolation of females prior to parturition reduces neonatal mortality in Galago

Eighteen years of birth records for three species of Galago at the Duke University Primate Center were examined to determine the effects of isolation of pregnant females on neonatal mortality rates. Isolation significantly decreased neonatal mortality rates in all three species over neonatal mortality rates in infants born to unisolated females. The frequency of cannibalism of infants did not differ between isolated and unisolated females. Secondary sex ratio differed significantly from 1:1 for all three species, but higher mortality in males in the first 10 days of life resulted in sex ratios that did not differ from 1:1 in G. garnettii and G. crassicaudatus.     

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.     

Differentiation of vocalizations in bushbabies (Galaginae, Prosimiae, Primates) and the significance for assessing phylogenetic relationships

A comparative bioacustic analysis of vocalizations of the prosimian subfamily Galaginae revealed that morphologically similar sibling taxa within the main groups of the lesser galagos and the greater galagos can be reliably identified phenotypically on the basis of the acoustic structure of their loud call or advertisement call. Results confirm the separation of two distinct species of greater galagos, Galago crassicaudatus and Galago garnettii, and strongly suggest the discrimination of three distinct species from the senegalensis lesser bushbaby group, Galago senegalensis, Galago moholi and Galao zanzibaricus. An investigation of the ontogenetic development of the loud call indicated that it is derived from the infant's isolation call, displaying in all studied bushbaby taxa a fairly similar acoustic pattern. Shared acoustic characters of the loud call among the different taxa as well as the infant's isolation call were used to propose a hypothesis about the phylogenetic affinities in bushbabies. The results seem to be supported by recent fossil records.     

Development of olfactory marker protein and N-CAM expression in chemosensory systems of the opossum,Monodelphis domestica

Using olfactory marker protein (OMP) and neural cell adhesion molecule (N-CAM) immunohistochemistry, the present study describes development of olfactory and vomeronasal systems in pre- and postnatal opossums, Monodelphis domestica. As revealed by OMP expression, development of the main olfactory system precedes that of the vomeronasal system by 1–2 weeks. OMP is expressed throughout (homogeneously) the nerve and glomerular layers of the main (MOB) but is expressed more strongly (heterogeneously) in the anterior as compared to the posterior accessory (AOB) olfactory bulb. N-CAM expression is homogeneous in both MOB and AOB. The heterogeneity in the AOB is developmentally regulated, since in the 30-day-old AOB the expression of OMP is homogeneous, becoming heterogeneous during the second month of life. Maximal expression of N-CAM precedes maximal expression of OMP in the VNS by about 2 weeks. From 7 to 21 days of age only, there is a small population of OMP-positive, N-CAM-negative olfactory and vomeronasal axon terminals that penetrate deep into the brain parenchyma, overgrowing their normal targets in the MOB and AOB, respectively. J. Morphol. 234:109–129, 1997. © 1997 Wiley-Liss, Inc.     

Syringeal anatomy and allometry in murres (Alcidae: Uria)

Species and sexual differences in vocalizations and the vocal tract are widespread. We studied the vocal tract of Common (Uria aalge) and Thick-billed (U. lomvia) Murres. We predicted anatomical or allometric differences in adults between species and sexes due to vocal differences related to social behavior (Common Murres nest at higher densities; males are the more aggressive sex, etc.). The vocal tract was anatomically simple and similar between species and sexes. The trachea was mainly cartilage, but the posteriormost 4–6 tracheal rings were calcified and fused as the tympanum, as part of the syrinx. The syrinx included the (unfused) first bronchial semirings, which were enlarged and calcified beneath the insertion of M. tracheolateralis. Weak bilateral asymmetry (right side larger) occurred in widths of M. tracheolateralis and M. sternotrachealis. The trachea was ∼10% longer in Common Murres; the tympanum and M. sternotrachealis width were relatively larger in Thick-billed Murres. Vocal-tract morphology and size did not differ between the sexes in either species. In allometric analyses on adults, isometry characterized (1) tympanum size in relation to head + bill length, and to humerus length (respectively), in Thick-billed Murres, and (2) M. sternotrachealis width in relation to tarsometatarsal length in both species. Comparative field studies on species and sexes are needed to clarify the functional significance of our findings in relation to vocalization.     

The vomeronasal organ and associated structures of the fetal African elephant, Loxodonta africana (Proboscidea, Elephantidae)

The vomeronasal organ (VNO) is a chemosensory structure of the nasal septum found in most tetrapods. Although potential behavioural correlates of VNO function have been shown in two of the three elephant species, its morphology in Loxodonta africana has not been studied. The development of the VNO and its associated structures in the African elephant are described in detail using serially sectioned material from fetal stages. The results show that many components of the VNO complex (e.g. neuroepithelium, receptor‐free epithelium, vomeronasal nerve, paravomeronasal ganglia, blood vessels, vomeronasal cartilage) are well developed even in a 154‐day‐old fetus, in which the VNO opens directly into the oral cavity with only a minute duct present. However, the vomeronasal glands and their ducts associated with the VNO were developed only in the 210‐day‐old fetus. Notably, in this fetus, the vomeronasal–nasopalatine duct system had acquired a pathway similar to that described in the adult Asian elephant; the VNOs open into the oral cavity via the large palatal parts of the nasopalatine ducts, which are lined by a stratified squamous epithelium. The paired palatal ducts initially coursed anteriorly at an angle of 45° from the oral recess and/or the oral cavity mucosa, and merged into the vomeronasal duct. This study confirms the unique characteristics of the elephant VNO, such as its large size, the folded epithelium of the VNO tube, and the dorsomedial position of the neuroepithelium. The palatal position and exclusive communication of the VNO with the oral cavity, as well as the partial reduction of the nasopalatine duct, might be related to the unique elephantid craniofacial morphogenesis, especially the enormous growth of the tusk region, and can be seen as autapomorphies.     

Milk composition reflects pattern of material care in prosimian primates

Hypothesized relationships between milk composition and life history traits were examined by analyzing mid-lactation milks of seven lemurs (Eulemur fulvus, E. macaco, E. rubriventer, E. mongoz, Varecia variegata, Hapalemur griseus, Lemur catta), three bushbabies (Otolemur crassicaudatus, O. garnettii, Galago moholi), and two lorises (Nycticebus coucang, Loris tardigradus); partial data were also obtained for the lemuroid Cheirogaleus medius. There were significant differences in milk composition among species within either Eulemur or Otolemur, but the four genera for which multiple samples were available (Eulemur, Varecia, Otolemur, and Nycticebus) exhibited large composition differences. Eulemur milk was, on average, very dilute (9.9% dry matter) and low in energy (0.49 kcal/g). These milks contained 0.9% fat, 1.2% protein, and 8.4% carbohydrate on a fresh weight basis. Protein energy comprised only about 15% of total milk energy. Varecia had significantly higher dry matter (13.5%), fat (3.2%), protein (4.2%), gross energy (0.80 kcal/g), and protein energy: total energy ratio (28%) than Eulemur. Milks of the lorisoid genera Otolemur and Nycticebus were very similar, and both had significantly higher dry matter (18.3, 16.3%), fat (7.6, 7.0%), and gross energy concentration (1.27, 1.13 kcal/g) than either lemuroid genus. Otolemur milk was higher in protein than Nycticebus milk. We conclude that lorises, bushbabies, and perhaps cheirogaleids produce relatively rich, energy-dense milks in comparison with anthropoid primates. However, dilute milks appear to be uniformly found among species of Eulemur and perhaps in Lemur catta. The milk of Varecia (and perhaps Hapalemur) is intermediate in composition. Differences in milk composition among prosimians may be related to differences in maternal care: prosimians that carry their young during lactation produce more dilute milks than do species which leave their young unattended for prolonged periods. When looking at primates as a whole, however, the picture is somewhat less clear, since the milks of some “parkers” like Varecia do closely resemble those of large anthropoid primates who carry their young. Am. J. Primatol. 41:195–211, 1997. © 1997 Wiley-Liss, Inc.     

性别、季节和体型大小对吐鲁番沙虎巢域的影响

2008年6月份至2009年5月份对吐鲁番沙虎的巢域进行调查:2008年6月份至2008年8月份为繁殖季节(RS),2008年9月份至2009年5月份(冬眠期除外)为非繁殖季节(NRS)。利用截趾标志重捕法研究吐鲁番沙虎的巢域,共标记283只吐鲁番沙虎,累计繁殖季节24只,非繁殖季节43只重捕超过3次(其中13只个体在繁殖季节和非繁殖季节均够3次以上捕捉次数,为重复个体),可以用于计算个体巢域面积数据。利用软件MapGis计算最小凸多边形法(MCP)巢域面积,并分析性别、体型大小、季节等因素对巢域的影响。结果表明:吐鲁番沙虎非繁殖季节雄性、雌性与幼体各组间的巢域面积差异均显著,繁殖季节巢域面积差异不显著;雌雄个体不同季节或全年合并比较巢域面积差异性均不显著;非繁殖季节面积与吻肛长(SVL)显著相关、全年成体组的巢域面积与吻肛长显著相关;成体巢域面积季节差异显著(U=41,P=0.046),幼体则没有季节差异(U=159,P=0.537)。因而,吐鲁番沙虎的巢域大小受性别因素影响不大,体型大小对巢域面积有显著影响,由于繁殖、食物资源等的季节变化是影响吐鲁番沙虎巢域最重要的因素。     

Species differences in substrate use and diet between sympatric galagos in two Kenyan coastal forests

Two sympatric species of galago,Galago zanzibaricus andG. garnettii, differed significantly in diet (judged mainly by fecal analysis) and substrate use at each of two coastal forest sites in Kenya. The larger-bodiedG. garnettii tended to use branches of a bigger diameter and to be found higher in the canopy than the smallerG. zanzibaricus. The latter species spent most of its time in the thick undergrowth. The larger species ate more fruit than the smaller. There were differences between the species in the type of invertebrate prey remains found in the feces. It is suggested that the differences in habitat use were influenced by the different body sizes of the two galagos, which so enabled these closely related species to coexist. Order of authorship is alphabetical.     

Separation possibilities and genital measurement variations in two cryptic species of European pierid butterflies,Leptidea juvernica Williams, 1946 and L. sinapis (Linnaeus, 1758)

Of the three recently separated cryptic butterfly species of the Leptidea sinapis complex, the two species L. sinapis and L. juvernica occur sympatrically and syntopically in central Europe. As the separation of these species requires genital or genetic characters, their correct identification is a fundamental problem to be solved prior to any biological and distributional studies. In the present study a morphometric approach was applied to test for separation possibilities and to examine genital measurement variations based on large population samples of the two species (347 females and 636 males). Butterflies were collected at 456 localities distributed across Poland. Specimens of both sexes could be separated using either the shape or the length of the antrum bursae (females), phallus and saccus (males). Intraspecific seasonal differences in these measurements may affect the discrimination of males only. Genital characters were significantly larger in butterflies of the spring brood than in those of the second brood. In females, antrum bursae length ranges were separated by an interval, allowing for the exact identification of each specimen. In males, no such intervals were found between phallus and saccus length ranges, which slightly overlapped. Discriminant analysis resulted in 100% of males correctly classified. The present study confirmed the validity of interspecific differences in the shape of phallus and saccus for species identification without measurements. The length of these two characters discriminates the two species when separation is limited to specimens from the same generations. As an alternative for discriminant analysis, either phallus length and vinculum width or phallus length and the ratio of the ventral edge of genital capsule length and saccus length (VEL/SL ratio) is recommended for the practical separation of the two species. The differences in the shape and size of the copulatory organs of the two species seem to indicate reproductive isolation due to mechanical incompatibility of their genitals.     

An Appraisal of Life History Features of Kiefferulus calligaster (Kieffer, 1911) (Diptera: Chironomidae) from Kolkata,India

Life history parameters of the freshwater chironomid species Kiefferulus calligaster (Kieffer, 1911) were investigated under laboratory conditions, with the use of larval development time and wing length as key features. An index of fitness was derived using these two parameters to represent the fitness of adults as a function of the larval development. Survivorship, deduced from the data on the mortality of larval stages, was related to developmental time as—(survivorship, l_x) y = 1.16 ? 0.04 × (days). The larval development time varied between males and females with a minimum of 8 and a maximum of 12 days from first instar larva to eclosion of imagine. The average wing length of adult females was larger than males (3.9 mm ± 0.03 S.E. vs. 3.36 mm ± 0.02 S.E.), for both early and late emerging individuals. The degree of dimorphism between the sexes was prominent for wing length and larval development time. The index of fitness for the early and late emerging adults differed significantly (P < 0.05) in both the sexes.