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.     

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.     

Hematology, serum chemistry values, and rectal temperatures of adult greater galagos (Galago garnetti and G. crassicaudatus)

Hematological and serum chemistry values, as well as rectal temperatures, were obtained from greater galagos (Galago garnettii and G. crassicaudatus), in order to establish normative values. No species or sex differences were found for four hematological parameters and 15 serum chemistry parameters. Species differences were seen in phosphate, magnesium, cholesterol, alkaline phosphate, G-glutamyl transferase, mean corpuscular volume and leucocyte, neutrophil, and lymphocyte number. Significant sex differences were observed in glucose, hemoglobin, and hematocrit values. Species and sex differences were seen in chloride and erythrocyte number.     

Effects of animate vs. inanimate stimuli on curiosity behavior in greater galago and slow loris

Captive prosimians housed in a semi-natural laboratory environment were exposed to novel stimuli, both animate (cat, snake) and inanimate (empty box). Observations were made by means of closed circuit TV. Animate stimuli elicited considerably more interest (about 42% of a test session) than the inanimate stimulus (about 12% of the test session). The more vulnerable species, the slow loris, displayed as much curiosity as the less vulnerable species, the greater galago; but, in the presence of live stimuli, lorises were more cautious. They moved slowly and silently, employed contact as a mode of investigation significantly less often than galagos, spent significantly less time than galagos investigating from within 3 ft, took significantly longer than galagos to make their first approaches to within the 3 ft zone, and, once there, stayed for significantly shorter visits than galagos. The data demonstrate: (1) that prosimians are highly responsive to novelty if the stimuli are appropriate; and (2) that it is important to take into account not just whether animals investigate but also how they investigate.     

Chromosomal evolution within the family Estrildidae (Aves) III. The Estrildae (waxbill finches)

The C- and G-banded karyotypes of five species of waxbill finches belonging to the Estrildidae were examined. Extensive chromosomal variation including inversions and fissions was found to differentiate the species, showing the waxbills to be the most chromosomally diverse group of estrildid finches. None of the variation, however, matched that recorded in related species of Pytilia (Christidis, 1983). By comparing the G-banded karyotypes of species belonging to all three estrildid lineages it was possible to reconstruct the presumed ancestral karyotype for the Estrildidae as a whole. This was found to approximate the karyotype of the Australian peophilid species, Poephila guttata most closely. From it chromosomal relationships within the three estrildid lineages, Poephilae, Lonchurae, Estrildae, can be determined.     

云南兔(Lepus comus)的染色体研究

本文以染色体分带技术(G-,C-带和银染色),对云南兔(Lepus comus)的核型进行了观察分析。结果表明t：2n=48,NF=88。常染色体由6对中着丝粒染色体, 5对亚中着丝粒染色体,10对亚端着丝粒染色体和2对端着丝粒染色体组成;X染色体为亚中着丝粒染色体。除少数几对染色体(Nos．9,14,16,20)着丝粒C-带呈阴性外,其余均呈阳性或浅染,但异染色质的含量差异较大。Ag-NORs的数目为5-6个,分别位于Nos．1 7,19,21。通过与兔科其它属种核型的比较,对其染色体进化作了讨论。     

Comparative penile morphology of East African galagos of the genus Galagoides (family Galagidae): implications for taxonomy

Recent studies on the comparative penile morphology of galagos have revealed complex patterns that vary among both genera and species, and help with species identification. So far the penile morphologies of 14 galago species have been described and an identification key has been proposed. The present study extends and revises previous work. Wild Galagoides cocos, G. granti, G. zanzibaricus and G. rondoensis from the Eastern Arc Mountains and coastal forests of Tanzania and Kenya were live-trapped, and one species (G. orinus) was examined using a museum specimen. Penile morphology was photographed, traced, and described qualitatively. All of the mature males had penile spines. Spines were absent or indistinct in immature males of all of the species. The penile morphologies of G. cocos, G. granti, and G. zanzibaricus are similar in their overall distribution and type of spines, but differ in the shape of the glans penis. Both G. orinus and G. rondoensis have divergent penile morphologies compared to all other galagos, and are probably phylogenetically distinct. The results support other recent morphological and behavioral studies that consider these five galagos to be distinct species. The evolution of the baculum and spines is thought to be linked to sexual selection in multimale mating systems, but the mechanisms involved are not entirely understood. Adult male G. rondoensis appear to display the penile spines by exposing the distal section of the penis.     

Body weight: its relation to tissue composition, segment distribution, and motor function. I. Interspecific comparisons.

The composition of skin, muscle, and bone and their distribution throughout the body are compared for "advanced" or "specialized" species (Alouatta, Macaca, Canis, Felis, Lepus); smaller, more closely related species (Tupaia and the Lorisidae); and several species within the same ecosystem (Barro Colorado Island, Panama). Among the most significant variables, the skin of sloths, howlers and macaques constitutes more than 12% of body weight, whereas greyhound skin is 5% of weight; sloth and howler muscle are 25% of weight, macaque muscle about 40% of weight, greyhound and agouti muscle over 50% of weight. In tree shrews and galagos muscle is heavier (35%) than in pottos and slow lorises (below 28%), but bone and skin are lighter. All species differ in the segmental distribution of weight. Cats have light tails, light feet and heavy thighs, whereas howlers have heavy tails, heavy feet, and light thighs. The galagos have heavy hindlimbs and tails, the pottos and lorises have reduced tails and approximately equal fore- and hindlimbs. Convergences in segment pattern (sloths with pottos and lorises, marmosets with tree shrews, owl monkeys with galagos, cebus with macaques) as well as divergences are documented. All weight-of tissue and weight-of-segment variables are correlated directly with locomotor adaptation.     

Sexual selection,genital morphology,and copulatory behavior in male Galagos

Variations in penile morphology among galago species are pronounced and complex. Comparative studies of galagos and other primate species show that elongation of the baculum (os penis)is associated with copulatory patterns involving a prolonged period of intromission. The enlarged penile “spines” of male galagos may be important in maintaining a genital “lock” during copulation. In primate species where females mate with a number of partners, sexual selection may have favored the rapid evolution of such features of penile morphology and masculine copulatory behavior. It is suggested that evolution of complex penile morphologies in galagos has been influenced by sexual selection and that such morphological variations are extremely useful in taxonomic studies.     

臭鼩染色体的研究

本文采用骨髓染色体制片法,对捕自我国浙江萧山市的臭鼩进行了组型、G-带、C-带和核仁组织区银染的观察分析。结果表明,我国臭鼩染色体数目为2n=40,组型为8(m)+2(sm)+10(st)+18(t),性染色体为,(?):X(m或sm),Y(m或sm);♀:XX(m或sm)。G-带较为丰富,每一对染色体都有其特定的带型,较易于辨别与配对。在C-带方前,4对中间着丝粒染色体与5对亚端着丝粒染色体均具有不同程度的着丝粒带,1对亚中着丝粒染色体与9对端着丝粒染色体缺乏C-带物质,性染色体具丰富的远端带及中间带.银染的结果显示,第5、12和13对染色体具银染物质。     

Mother-infant cannibalism in thick-tailed bushbabies (Galago crassicaudatus umbrosus)

Mother-infant cannibalism in species of galagos as in several other species of non-human primates is a common phenomenon. In non-human primates kept in laboratory conditions many of the observed cases of cannibalism were not associated with starvation and with infanticide. Cannibalism in galagos was observed in at least five different species. In several laboratories, like the Duke University Primate Center, the frequencies of cannibalism in galagos species in captivity have been reduced by the isolation of pregnant female before the parturition from her companions, especially adult males. At the Primate Behaviour Research Group (University of Witwatersrand, Johannesburg, South Africa) three cases of cannibalism were observed in twoGalago crassicaudatus umbrosus' mothers before the end of the third day after birth. To understand the reasons why the mothers cannibalize their own infants, it was decided to analyze the mother-infant interaction which preceded the death of the infants. InGalago crassicaudatus twin and triplet births are very common. In one of our two observed mothers who cannibalized their infants, there was one triplet birth. In this study the two pregnant females were isolated in two separated cages ten days before giving birth. One of the two observed mothers gave birth to triplets, the other had a single birth. At the end all four infants were cannibalized. In all cases the cannibalization started after the infant's death. The infants' deaths were caused by lack of maternal care which caused them to starve. The results show that the triplet's mother differed in some of her behaviour towards her infants. In one of the cannibalized infants behaviours such asNipple andMother Passive Prevent were never seen before death. In our cases it seems that cannibalism was probably due to the laboratory condition that for generation to generation caused stress to the mothers and changed their maternal behaviours, which lead to the death of the offspring which after death became a consumable resource.     

Comparative analysis of karyotypes in European shrew species. I. The sibling species Sorex araneus and S. gemellus: Q-bands, G-bands, and position of NORs

The karyotypes of two closely related species of the genus Sorex (Mammalia, Insectivora) were compared with each other by G- and Q-banding techniques and by Ag-AS staining (GOODPASTURE and BLOOM, 1975). By comparing the G-banded karyotypes, it could be ascertained that the basic differences in karyotype between the two species lie in three pericentric inversions, three paracentric inversions, and one reciprocal translocation. This is in near agreement with FORD and HAMERTON (1970), who assumed that both species differ by three pericentric inversions and one tandem translocation. Furthermore, the karyotype of S. araneus (race C) presented by HALKKA et al. (1974) has been compared with the S. araneus of the present report. Considering the species with respect to karyotypic evolution, it is supposed that S. araneus and S. gemellus derive from a common ancestor.     

Masseter electromyography during chewing in ring-tailed lemurs (Lemur catta)

We examined masseter recruitment and firing patterns during chewing in four adult ring-tailed lemurs (Lemur catta), using electromyography (EMG). During chewing of tougher foods, the working-side superficial masseter tends to show, on average, 1.7 times more scaled EMG activity than the balancing-side superficial masseter. The working-side deep masseter exhibits, on average, 2.4 times the scaled EMG activity of the balancing-side deep masseter. The relatively larger activity in the working-side muscles suggests that ring-tailed lemurs recruit relatively less force from their balancing-side muscles during chewing. The superficial masseter working-to-balancing-side (W/B) ratio for lemurs overlaps with W/B ratios from anthropoid primates. In contrast, the lemur W/B ratio for the deep masseter is more similar to that of greater galagos, while both are significantly larger than W/B ratios of anthropoids. Because ring-tailed lemurs have unfused and hence presumably weaker symphyses, these data are consistent with the symphyseal fusion-muscle recruitment hypothesis stating that symphyseal fusion in anthropoids provides increased strength for resisting forces created by the balancing-side jaw muscles during chewing. Among the masseter muscles of ring-tailed lemurs, the working-side deep masseter peaks first on average, followed in succession by the balancing-side deep masseter, balancing-side superficial masseter, and finally the working-side superficial masseter. Ring-tailed lemurs are similar to greater galagos in that their balancing-side deep masseter peaks well before their working-side superficial masseter. We see the opposite pattern in anthropoids, where the balancing-side deep masseter peaks, on average, after the working-side superficial masseter. This late activity of the balancing-side deep masseter in anthropoids is linked to lateral-transverse bending, or wishboning, of their mandibular symphyses. Subsequently, the stresses incurred during wishboning are hypothesized to be a proximate reason for strengthening, and hence fusion, of the anthropoid symphysis. Thus, the absence of this muscle-firing pattern in ring-tailed lemurs with their weaker, unfused symphyses provides further correlational support for the symphyseal fusion late-acting balancing-side deep masseter hypothesis linking wishboning and symphyseal strengthening in anthropoids. The early peak activity of the working-side deep masseter in ring-tailed lemurs is unlike galagos and most similar to the pattern seen in macaques and baboons. We hypothesize that this early activity of the working-side deep masseter moves the lower jaw both laterally toward the working side and vertically upward, to position it for the upcoming power stroke. From an evolutionary perspective, the differences in peak firing times for the working-side deep masseter between ring-tailed lemurs and greater galagos indicate that deep masseter firing patterns are not conserved among strepsirrhines.     

The chromosomes of Nycticebus coucang (Boddaert, 1785) (Primates: Prosimii)

The high-quality karyotype of a specimen of Nycticebus coucang is described and illustrated. The X chromosome is found to be indistinguishable from that of the greater galagos, and may represent a synapomorphic trait. The Y chromosome is a medium to small submetacentric (3.2% TCL) and constitutes one of the larger Y chromosomes known in primates. N. coucang is found to have multiple NOR-bearing chromosomes in contrast to the single pair found in galagine and catarrhine monkeys. Since a single NOR-bearing pair is often considered ancestral for primates, this new finding may have important implications for the evolution of these cistrons. One of the chromosomal polymorphisms in this specimen is a pericentric inversion, involving a NOR-bearing autosomal pair (no. 6), that alters the position of the active site. Further, homologues 2p differ by aparacentric inversion. These results confirm that lorisiforms are characterized by considerable chromosomal polymorphism.     

Galago locomotion in Kibale National Park, Uganda

Very few locomotor studies have been conducted on galagos. This is surprising given their interesting anatomy and ecology, as well as their increasing species diversity. In this study we investigated locomotion and postures in two sympatric galagos species (Galagoides thomasi and Galago matschiei) living in Kibale National Park, Uganda. G. thomasi uses arboreal quadrupedalism and leaping, while G. matschiei is more leaping-oriented. Both species utilize small oblique branches in the mid-canopy. These similarities in substrate use are most likely due to the similar body sizes and anatomies of the two species, as well as to the structure and availability of trees in Kibale National Park. Lastly, we compare the locomotor patterns of G. thomasi and G. matschiei with those observed in the few other quantitative locomotor studies available for galagos.     

Chromosome banding patterns of the aye-aye,Daubentonia madagascariensis (primates,Daubentoniidae)

The aye-aye (Daubentonia madagascariensis) is an endangered prosimian primate found only on the island of Madagascar. It is the only extant representative of its family Daubentoniidae. The phylogenetic relationship of this species with other prosimians is unclear. Because a G-banded karyotype forDaubentonia has not previously been reported, blood for preparation of lymphocyte cultures was obtained from one of the four aye-ayes in captivity in the United States. The diploid chromosome number was 30. The karyotype consisted of 14 autosomal metacentrics, 10 autosomal submetacentrics, and 4 autosomal acrocentrics. The similarities between the G-banded chromosomes ofDaubentonia and those ofPropithecus, a member of the Indriidae, support the notion thatDaubentonia has a closer relationship with the Indriidae than with other Lemuriformes or the lorisoids.     

Effects of ovariectomy and estradiol replacement therapy upon the sexual and aggressive behavior of the greater galago (Galago crassicaudatus crassicaudatus)

The effects of ovariectomy and estradiol treatment upon sexual and aggressive behavior were studied in a prosimian primate, the greater galago. Ovariectomized galagos were sexually unreceptive and frequently aggressive, but retained their sexual attractiveness to males. When females were treated with estradiol monobenzoate, however, their aggression and refusals of males' mounting attempts decreased markedly. Although males mounted these females, they usually failed to copulate, possibly because the females did not perform certain postural adjustments which assist males to intromit. Estradiol benzoate alone, even in large doses, does not fully restore patterns of mating behavior in ovariectomized female greater galagos. These observations on a prosimian primate are in striking contrast to the results of similar work on Old World monkeys and chimpanzees.     

Tertiary sex ratios in wild Galago population (Mammalia: Primates)

Previous studies have proposed that species within the genus Galago exhibit a secondary sex ratio that is strongly skewed (approximately 70%) in favour of males. It has further been suggested that this sex ratio is maintained into adulthood. We present data obtained from 669 adult galagos, representing eight wild populations and five species, to indicate that the sex ratio among sexually mature individuals is close to parity. If the observations regarding sex ratios at birth are correct, this implies that juvenile/sub-adult males suffer significantly higher mortalities than do their female counterparts.     

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.