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.     

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.     

Reproductive patterns in galagos (Galago zanzibaricus and Galago garnettii) in relation to climatic variability

The reproductive patterns (birth seasonality, litter size, litters per year) of two sympatric species of galago (Galago zanzibaricus and G. garnettii) were studied in a coastal forest in Kenya for a two-year period. Trap-retrap and radio tracking methods were employed. G. zanzibaricus has one infant twice per year; G. garnettii has one infant once per year. Both species are seasonal breeders. These East African galagos are intermediate in reproductive patterns when compared with galagos from South African woodland (G. senegalensis moholi and G. crassicaudatus umbrosis) and West African rainforest (G. alleni and G. demidovii). Climatic patterns (total annual rainfall, seasonal variability of rainfall, variability in total annual rainfall, and annual temperature variability) are also compared for the three regions. Climatically, East Africa is intermediate between West and South Africa in total annual rainfall and in seasonality of rainfall, but not in year-to-year variability in rainfall. East Africa shows the highest variability in annual rainfall. South Africa has the coldest dry seasons and highest variability in temperatures. The results of this study suggest that “r-selection” and “K-selection” do not provide adequate explanations of galago reproductive patterns.     

Reproductive and dietary parameters in wild greater galago populations

Information concerning habitat, body size, reproductive status, and diet was recorded from 348 greater galagos, captured at six different localities in Tanzania and southern Africa between 1953 and 1955. The localities extended from Pemba Island in the north to Chikwawa, Malawi, in the south and varied broadly in the same order in degree of climatic aridity— from well-watered clove and coconut plantations to seasonally very dry woodland. Animals from the three northern localities fell within the geographic range of Galago garnettii,while the rest were assumed to be G. crassicaudatus.Statistical analysis of body size parameters confirmed this allocation. Data on fetal occurrence, vaginal and labial condition, and lactation indicate a restricted breeding season for both species, with peak proportions in estrus in August in G. garnettiiand in May-July in G. crassicaudatus.Gut content data indicate a variety of foods, with a preponderance in the northern localities of soft fruit such as mango, pawpaw, and coconut pulp; gum was a major carbohydrate source in the southernmost localities. Contrary to expectations, tooth damage, in the form of both loss and breakage, was much more prevalent in G. garnettiithan in G. crassicaudatus.The teeth most commonly lost were the upper incisors— perhaps because of the high acid and sugar content of a frugivorous diet. The high incidence of breakage of the lower incisors and upper canines indicates the inclusion of hard-shelled food sources in the diet of G. garnettii.     

Social organization of Galagos in Kenyan coastal forests: I. Galago zanzibaricus

The social organization of Galago zanzibaricus 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. Associations in sleeping groups indicated social ties between individuals. Patterns of range use were established by radio-tracking focal individuals. Adult males generally had nonoverlapping ranges, shared with one or two females and their offspring with which the male regularly slept. Young females remained longer in their natal ranges than males and reproduced within their natal ranges. Range size and distance traveled per night were similar between the sexes; both sexes probably defended territories. None of the other galago (bushbaby) species studied to date show the degree of close male-female association found in G. zanzibaricus. This study thus extends the variety of social organization documented in nocturnal prosimians.     

Seasonal variation in the diet of South African Galagos

The feeding behavior and diet of two species of bushbaby, Galago senegalensisand Galago crassicaudatus,in South Africa were examined in relation to seasonal changes in climate. The populations studied were allopatric, but both fed predominantly on Acaciagum and invertebrates. Data on their diet were collected by direct observation of a radiocollared female of each species and by analysis of fecal samples. Differences in diet were found between the species within seasons and within species between the seasons. Both spent more time gum-feeding in winter than in summer. Prey size and type differed between the species in summer but converged in winter. The quantity of insects taken was fairly similar between seasons for G. senegalensisbut dropped considerably for G. crassicaudatusin winter. These observations are interpreted in terms of interspecific differences in feeding strategies, which are considered to be a function of the difference in body size of the two galago species.     

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 taxonomic status and distribution of Bushbabies in Malawi with emphasis on the significance of vocalizations

The debated identity of a small forest bushbaby in Malawi is resolved by a short-term field study of the animals’ behavior. Locomotor styles, calling patterns, and the structure of advertising calls confirm that the species is Galago zanzibaricusrather than G. demidoffor G. thomasi.A detailed comparison of acoustic structure between the Malawi animals and G. zanzibaricusin Kenya demonstrates a degree of between-population variation, although the calls remain conservative in those parameters expected to aid recognition of conspecifics. Distribution records extend the known geographical range of G. zanzibaricusover most of the northern half of Malawi. Further studies are required to link the animals from this region with either of the previously recognized subspecies: G. z. zanzibaricusfrom East Africa or G. z. grantifrom southern Malawi, Mozambique, and Zimbabwe.     

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.     

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.     

Polytene chromosome maps in four species of tsetse flies Glossina austeni, G. pallidipes, G. morsitans morsitans and G. m. submorsitans (Diptera: Glossinidae): a comparative analysis

Photographic polytene chromosome maps from pupal trichogen cells of four tsetse species, Glossina austeni, G. pallidipes, G. morsitans morsitans and G. m. submorsitans were constructed and compared. The homology of chromosomal elements between the species was achieved by comparing banding patterns. The telomeric and subtelomeric chromosome regions were found to be identical in all species. The pericentromeric regions were found to be similar in the X chromosome and the left arm of L1 chromosome (L1L) but different in L2 chromosome and the right arm of L1 chromosome (L1R). The L2 chromosome differs by a pericentric inversion that is fixed in the three species, G. pallidipes, G. morsitans morsitans and G. m. submorsitans. Moreover, the two morsitans subspecies appeared to be homosequential and differ only by two paracentric inversions on XL and L2L arm. Although a degree of similarity was observed across the homologous chromosomes in the four species, the relative position of specific chromosome regions was different due to chromosome inversions established during their phylogeny. However, there are regions that show no apparent homology between the species, an observation that may be attributed to the considerable intra—chromosomal rearrangements that have occurred following the species divergence. The results of this comparative analysis support the current phylogenetic relationships of the genus Glossina.     

Interference competition in desert subterranean termites

We examined interspecific aggression between two subterranean termite species, Heterotermes aureus (Snyder) (Rhinotermitidae) and Gnathamitermes perplexus (Banks) (Termitidae). In laboratory tests with worker termites, neither species was the inherently superior fighter, but rather the outcome of interspecific encounters depended on the number of conspecifics. We then investigated patterns of resource use by these species during a 13-month period in the Sonoran Desert. Baits consisted of toilet-paper rolls, which have been shown to be a mutually acceptable food source. Analyses of foraging activity demonstrated that the two species did not forage independently of each other. Not only were the two species negatively associated spatially, but extended periods of temporal segregation were observed. G. perplexus took significantly longer to return to sites that it had simultaneously occupied with H. aureus than to sites that G. perplexus had occupied alone. The pattern of co-occurrence of these two species is consistent with the hypothesis that interspecific interference competition affects their spatial and temporal distribution.     

Molecular divergence in the Galapagos Islands—Baja California species pair,Gossypium klotzschianum andG. davidsonii (Malvaceae)

Molecular divergence betweenGossypium klotzschianum andG. davidsonii was studied. The former is endemic to five of the larger islands of the Galapagos, whileG. davidsonii is restricted to the southern half of Baja California, approximately 2 500 km distant. A substantial body of genetic and taxonomic data suggests that these two species are related as progenitor and derivative. Interspecific hybrids are fully fertile, with no evidence of F₂ breakdown and normal segregation of genetic markers. Allozyme analysis of 33 populations for 41 loci indicated that the allelic composition ofG. klotzschianum represents a subset ofG. davidsonii. Although genetic diversity is relatively restricted in both species, calculated measures demonstrate higher levels of genetic variability and greater population structuring inG. davidsonii than inG. klotzschianum. The interspecific genetic identity of 0.87 is typical for progenitor-derivative species pairs. Chloroplast DNAs were surveyed for variation with 25 restriction enzymes using hybridization probes that cover the entire chloroplast genome. No intraspecific and little interspecific variation was detected among 560 cpDNA restriction sites, representing sequence information for approximately 3200 nucleotides. Only 3 mutational differences distinguished the two species, resulting in a sequence divergence estimate of 0.09%. Divergence times were estimated from both the isozyme data and the cpDNA restriction site data. Although these estimates have several sources of error, both molecular data sets were congruent in suggesting that the two lineages diverged between 250000 and 700000 years ago. Accumulated evidence suggests that dispersal was from Baja California to the Galapagos Islands rather than the reverse, and most likely was mediated by trans-oceanic drift.G. klotzschianum may be the only species of the endemic Galapagos flora to have arisen from a northern Mexican progenitor.     

桉树杂交种对桉树枝瘿姬小蜂的抗性变异分析

植物种间杂交是一种普遍自然现象,杂交往往造成植物表型及生理变异,从而改变杂种抗虫性。与亲本种相比,杂种抗虫性可能增强或减弱,也有可能处于与亲本相似水平。初生、次生代谢物的质变与量变是引起杂种抗虫性变异的重要原因。近年来,桉树杂交育种已在世界范围内广泛应用并取得了显著成效,桉树杂交种间抗虫性表现参差不齐,因此,桉树是研究杂交种抗虫性变异机制的理想材料。以2个桉树杂交种巨细桉DH201-2、巨尾桉G9及桉树重要害虫桉树枝瘿姬小蜂为研究对象,比较了2个杂交种与其纯亲本种[(巨桉×细叶桉),(巨桉×尾叶桉)]间的抗虫性差异;同时,综合比较了品系间叶片性状(叶片厚度、含水率、比叶面积)、初生化合物(C、N、可溶性糖、可溶性蛋白)及次生化合物(总酚、单宁)差异,以研究桉树杂交种抗虫性变异的理化机制。结果表明:DH201-2感染桉树枝瘿姬小蜂的虫瘿数目显著高于其双亲本种,而G9上虫瘿数目显著低于其双亲本种。DH201-2与G9的叶片厚度与巨桉相近,而显著薄于另一亲本种。DH201-2叶片含水率显著高于细叶桉、与巨桉相近;G9叶片含水率则显著低于其双亲本种。相似的是,DH201-2和G9的比叶面积均显著高于其双亲本种。初生化合物方面,DH201-2叶片可溶性糖和可溶性蛋白含量均显著高于其亲本种,N含量则仅高于细叶桉;而G9叶片可溶性蛋白含量虽高于其双亲本种,可溶性糖含量则无显著差异,N含量显著低于其双亲本种。次生化合物方面,DH201-2叶片总酚和单宁含量显著低于其双亲本种,而G9则显著高于其双亲本种。因此,与其亲本种相比,DH201-2感虫性增加,而G9抗虫性增加;与桉树枝瘿姬小蜂发育相关的营养指标(如含水率、可溶性糖、N含量)及次生防御物质(如总酚、缩合单宁)在桉树杂交种组织内的含量差异影响了桉树杂交种对桉树枝瘿姬小蜂的抗性。在全球推行桉树杂交育种且桉树害虫数量逐年增加的大背景下,应加强对桉树杂交种抗虫性机制研究,为选育高抗品系及桉树产业可持续发展提供理论指导。     

Field water relations of three temperate vines

Summary A two year comparative field water relations study was conducted in central Maryland on three sympatric temperate lianas, Lonicera japonica, Vitis vulpina and Parthenocissus quinquefolia. Seasonal physiological activity was longer by approximately 9 weeks in the evergreen L. japonica, while peak rates of stem elongation were 4–10 fold higher in the two deciduous species. There were marked differences in vascular anatomy and water use patterns among the three species, however all three evidenced varying degrees of stomatal closure in response to decreasing soil water availability and increasing atmospheric evaporative demand. The range of leaf water potentials measured in these species was quite narrow in comparison to other temperate woody species. Two of the species showed no alterations in their tissue water release properties in response to decreased soil water availability, while V. vulpina showed a limited capacity in this regard. Most significant among the species differences in water relations were the conservative water use patterns of P. quinquefolia, and the midday maxima of transpirational water loss measured in L. japonica compared to the morning peaks in traspiration for the two deciduous species. The differences found in anatomy, leaf phenology, climbing mechanics, water relations and canopy development among these three sympatric vines implies a spatial and temporal partitioning of light and water resources and emphasizes the diversity of morphological-physiological suites of characters present among species co-occurring in the same macrohabitat.     

Ecological attributes recorded in stable isotope ratios of arboreal prosimian hair

Carbon and nitrogen stable isotope ratios were measured in hair samples from two species of Galago from Gedi Ruins National Monument in eastern Kenya and from Lepilemur leucopus from Beza Mahafaly Special Reserve in southern Madagascar. Forest structure was generally similar in the two areas but average rainfall was lower in Madagascar. Species average ¹³C values varied with feeding height in the forest canopy and with average rainfall level as expected from reported variation in plant ¹³C values. G. garnettii, which feeds higher in the forest canopy, had less negative ¹³C values than G. zanzibaricus, which spends more time below 5 m. L. leucopus, from a drought-afflicted forest, had less negative hair ¹³C values than the two galago species. The values within the Lepilemur sample showed a positive linear relation with percent dependence on a CAM tree species and with xeric conditions within the species reserve. Nitrogen stable isotope ratios varied with trophic level of feeding and with time spent feeding on leguminous plants. The insectivorous galagos had significantly more positive ¹⁵N values than the folivorous L. leucopus. Within the Lepilemur sample, ¹⁵N values varied inversely with the percent of feeding time spent on leguminous plants. The range of ¹⁵N and ¹³C values in each of the prosimian species is larger than reported for animals fed monotonous diets and for New World monkey species. The monkey species feed as groups of individuals whereas the prosimians have solitary feeding habits. The ranges in the prosimian species apparently reflect the greater variation in diet among individual prosimians compared to individual monkeys. The isotope data reported here are equivalent, on average, to those reported for other arboreal species from similar forest habitats and with similar dietary habits. This supports the use of such data for paleoecological reconstruction of forest and woodland systems and diet reconstruction of extinct primate populations and species. Received: 18 April 1997 / Accepted: 11 August 1997     

Comparative embryology of threeGentianaceae species from the Central Caucasus and the European Alps

A comparative investigation was carried out on the ovule and seed development of three mountain species ofGentianaceae, the perennial speciesGentiana pyrenaica, and the two short-lived monocarpic speciesGentianella caucasea andG. germanica. In all three species most embryological characters conform to those generally found in the family of theGentianaceae. In some features, however,G. pyrenaica and the twoGentianella species differ from each other. InG. pyrenaica the ovule is anatropous, the integument 8–10 layered and the three reduced antipodals degenerate soon after fertilization. In contrast,G. caucasea andG. germanica form a hemitropous ovule, a 4–5 layered integument and up to 16 antipodal cells by secondary multiplication. All three species exhibit differences in synchronization between embryogenesis and endosperm development. Functional relations between the antipodal structure and the dynamics of seed development of the investigated species are postulated.     

Contrasting reproductive parameters ingalago senegalensis braccatus andG. s. moholi

Reproductive parameters were compared between two subspecies of the lesser bushbaby, Galago senegalensis braccatusand G. s. moholi.Adult body weights for both male and female G. s. braccatuswere significantly higher than adult body weights of G. s. moholi,as were infant birth weights. The mean gestation length of G. s. braccatuswas significantly longer than that of G. s. moholi. G. s. moholihad a higher incidence of twinning than G. s. braccatus.Puberty in females occurred significantly later in G. s. braccatusthan in G. s. moholi.These differences are of such magnitude that hybridization of G. s. braccatuswith G. s. moholiwould be unlikely. These differences, if augmented by additional evidence, may warrant distinction of the two subspecies at the species level.     

Studies on the diversity of the distinct phylogenetic lineage encompassing Glomus claroideum and Glomus etunicatum

Morphological and molecular characters were analysed to investigate diversity within isolates of the Glomus claroideum/Glomus etunicatum species group in the genus Glomus. The inter- and intra-isolate sequence diversity of the large subunit (LSU) rRNA gene D2 region of eight isolates of G. claroideum and G. etunicatum was studied using PCR-single strand conformational polymorphism (SSCP)-sequencing. In addition, two isolates recently obtained from Southern China were included in the analysis to allow for a wider geographic screening. Single spore DNA isolation confirmed the magnitude of gene diversity found in multispore DNA extractions. An apparent overlap of spore morphological characters was found between G. claroideum and G. etunicatum in some isolates. Analysis of the sequence frequencies in all G. etunicatum and G. claroideum isolates (ten) showed that four LSU D2 sequences, representing 32.1% of the clones analysed for multispore extraction (564) were found to be common to both species, and those sequences were the most abundant in four of the ten isolates analysed. The frequency of these sequences ranged between 23.2% and 87.5% of the clones analysed in each isolate. The implications for the use of phenotypic characters to define species in arbuscular mycorrhizal fungi are discussed. The current position of G. claroideum/G.etunicatum in the taxonomy of the Glomeromycota is also discussed.     

Search for effective natural enemies of Tetranychus evansi in south and southeast Brazil

Tetranychus evansi Baker & Pritchard is an important pest of Solanaceae in several countries. Introduced accidentally to Africa, it presently occurs in many countries of that continent. In some of them, it is considered a key pest. The suspected area of origin of this mite is South America. The objective of the present study was to identify phytoseiid mites on solanaceous plants in association with T. evansi in south and southeast Brazil for introduction in the African continent for use in a classic biological control program. Almost 1,400 predatory mites of the family Phytoseiidae were collected, on 22 solanaceous species. The Amblyseiinae were the most diverse group in this study. Twenty-three of the species found belong to this subfamily, while only three belong to the Typhlodrominae and two to the Phytoseiinae. The most abundant and most frequent phytoseiid species were Phytoseius guianensis De Leon and Galendromus annectens (De Leon) of the Phytoseiinae and Typhlodrominae, respectively. The most frequent and abundant species of Amblyseiinae was Neoseiulus tunus (De Leon). Phytoseius guianensis and N. tunus were never found in association with T. evansi and G. annectens was found only once in association with it. Two factors suggested Phytoseiulus longipes Evans as the most promising predator found in this study. It could walk very well on tomato leaves infested by T. evansi, without being hampered by the profuse webbing produced by the prey and by the trichomes. In addition, several specimens of both sexes including eggs and nymphs of the predator were found associated with T. evansi on three different plant species and in two different periods of the year, when T. evansi was the only arthropod present on the leaves.