New species of the fleas Farhangia and Nestivalius,from endemic rodents in Sulawesi,Indonesia

The fleas (Siphonaptera: Pygiopsyllidae) Farhangia quattuordecimdentata sp. n. and Farhangia sedecimdentata sp. n. are described from pygmy squirrels (Prosciurillus spp.), and Nestivalius sulawesiensis sp. n. from murine rodents, all from Sulawesi, Indonesia. Both new species of Farhangia were collected in Central Sulawesi (Sulawesi Tengah); F. quattuordecimdentata sp. n. was recovered mainly from P. murinus, whereas F. sedecimdentata sp. n. was recovered mainly from P. leucomus. These new species are compared with the two previously described species of Farhangia: F. celebensis (Ewing) from P. murinus in North Sulawesi (Sulawesi Utara) and F. sciuri (Ewing) from the tree squirrel Callosciurus prevosti in Sabah (Borneo). Nestivalius sulawesiensis sp. n. was collected from six species of endemic murine rodents in both North and Central Sulawesi. It is compared to the morphologically similar N. pomerantzi (Traub) from Mindanao, which parasitizes murine hosts that are endemic to the Philippines.     

Tree diversity in primary forest and different land use systems in Central Sulawesi,Indonesia

We studied the tree communities in primary forest and three different land use systems (forest gardens, ca. 5-year-old secondary forests, cacao plantations) at 900–1200 m elevation in the environs of Lore Lindu National Park, Central Sulawesi. The primary forests had ca. 150 tree species 10 cm diameter at breast height (dbh) per hectare, which is unusually high for forests at this elevation in southeast Asia. Basal area in the primary forest was 140 m² ha^–1, one of the highest values ever recorded in tropical forests worldwide. Tree species richness declined gradually from primary forest to forest gardens, secondary forests, and cacao plantations. This decline was paralleled by shifts in tree family composition, with Lauraceae, Meliaceae, and Euphorbiaceae being predominant in primary forests, Euphorbiaceae, Rubiaceae and Myristicaeae dominating in the forest gardens and Euphorbiaceae, Urticaceae, and Ulmaceae in the secondary forests. Cacao plantations were composed almost exclusively of cacao trees and two species of legume shade trees. Forest gardens further differed from primary forests by a much lower density of understorey trees, while secondary forests had fewer species of commercial interest. Comparative studies of birds and butterflies demonstrated parallel declines of species richness, showing the importance of trees in structuring tropical forest habitats and in providing resources.     

The influence of emergent trees on rainfall distribution in a cacao agroforest (Sulawesi,Indonesia)

Emergent trees may have an influence on the volume and the spatial distribution of water input into agroforestry stands and may thus affect water availability for the main crops. Our goal was to analyze the influence of such trees on rainfall distribution in a cacao agroforest area in the rainforest margin zone of Central Sulawesi, Indonesia. The emergent trees studied belong to the species Bischofia javanica (Phyllanthaceae) and were 15 m high remnants from the natural forest. A set of 96 throughfall gauges was systematically distributed underneath canopies of cacao only, and underneath canopies of cacao plus emergent trees (cacao plus trees). From an earlier study we knew that stemflow can safely be estimated with less than 1% of gross precipitation (Pg).     

Rapid evolution of sessility in an endemic species flock of the freshwater bivalve Corbicula from ancient lakes on Sulawesi, Indonesia

The fauna of ancient lakes frequently contains taxa with highly derived morphologies that resulted from in situ radiation of lacustrine lineages with high antiquity. We employed a molecular mtDNA phylogeny to investigate this claim for corbiculid freshwater bivalves in two ancient lake systems on the Indonesian island Sulawesi. Among the otherwise mobile corbiculid species flock, only one taxon, Posostrea anomioides, in the ancient Lake Poso exhibits a unique habit, i.e. cementing one valve to the substrate. Our data show that Corbicula on Sulawesi is polyphyletic, with the endemic riverine taxa in terminal position, and the lacustrine species flock being paraphyletic. Surprisingly, Posostrea is not confirmed as a genus distinct from Corbicula and genetic distances suggest a rather recent origin from the only other corbiculid species endemic to Lake Poso, the non-cementing Corbicula possoensis. While the cementing anomioides, despite its unique behavioural and morphological characteristics, clusters together with non-sessile Corbicula species, the latter exhibit strong genetic distances in the absence of morphological disparity and fall into several genetically rather distinct clades. These findings suggest that developmental plasticity of animals in ancient lakes rather than the antiquity of lineages might account for the unique morphology of some species.     

Hybridization and population genetics of two macaque species in Sulawesi, Indonesia

This study investigates hybridization and population genetics of two species of macaque monkey in Sulawesi, Indonesia, using molecular markers from mitochondrial, autosomal, and Y-chromosome DNA. Hybridization is the interbreeding of individuals from different parental taxa that are distinguishable by one or more heritable characteristics. Because hybridization can affect population structure of the parental taxa, it is an important consideration for conservation management. On the Indonesian island of Sulawesi an explosive diversification of macaques has occurred; seven of 19 species in the genus Macaca live on this island. The contact zone of the subjects of this study, M. maura and M. tonkeana, is located at the base of the southwestern peninsula of Sulawesi. Land conversion in Sulawesi is occurring at an alarming pace; currently two species of Sulawesi macaque, one of which is M. maura, are classified as endangered species. Results of this study indicate that hybridization among M. maura and M. tonkeana has led to different distributions of molecular variation in mitochondrial DNA and nuclear DNA in the contact zone; mitochondrial DNA shows a sharp transition from M. maura to M. tonkeana haplotypes, but nuclear DNA from the parental taxa is homogenized in a narrow hybrid zone. Similarly, within M. maura divergent mitochondrial DNA haplotypes are geographically structured but population subdivision in the nuclear genome is low or absent. In M. tonkeana, mitochondrial DNA haplotypes are geographically structured and a high level of nuclear DNA population subdivision is present in this species. These results are largely consistent with a macaque behavioral paradigm of female philopatry and obligate male dispersal, suggest that introgression between M. maura and M. tonkeana is restricted to the hybrid zone, and delineate one conservation management unit in M. maura and at least two in M. tonkeana.     

Elevational patterns of species richness and density of rattan palms (Arecaceae: Calamoideae) in Central Sulawesi, Indonesia

We studied species richness and density of rattan palms in 50 plots of 10 × 100 m² each between 250 m and 2420 m in eight sites in Lore Lindu National Park (LLNP), Central Sulawesi. Rattans were observed in all sample sites, representing three genera and 34 species. The elevational patterns for species richness and density were humped-shaped with maxima around 1000 m. Polynomial models of second order explained 59 and 32% of species richness and density with the factor elevation, respectively. A majority of rattan species (65%) overlapped between 1000 and 1100 m elevation, while a pronounced change in the rattan flora occurred above 1100 m. Commercially important rattan species (Calamus zollingeri, C. ornatus var. celebicus, Daemonorops macroptera) were not observed above 1250 m. The change of species assemblage was significantly related to elevation (56%), followed by geographical distance (47%) and precipitation (40%). Less than 10% of LLNP is lowland forests, much of which is threatened by agricultural intensification. In contrast, montane forests are well represented in the park and high elevation forests are not subject to agricultural conversion or intensive harvesting of rattan and other forest products.     

Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.     

Population densities of Sulawesi crested black macaques (Macaca nigra) on Bacan and Sulawesi,Indonesia: Effects of habitat disturbance and hunting

Population surveys of Sulawesi crested black macaques (Macaca nigra) were conducted on the Indonesian islands of Sulawesi and Bacan in 1992–1994 to assess the status of natural populations and determine habitat and anthropogenic factors affecting their population densities. We surveyed five sites for primates, including undisturbed and disturbed habitats. Data were collected on habitat structure and composition at two undisturbed and one disturbed forest site in which the primates were surveyed. The highest density of macaques was found in primary forest at Gunung Sibela Nature Reserve on Bacan (170.3 individuals/km²). Population density in logged forest on Bacan was high but significantly less than primary forest (133.4 individuals/km²). The high density of crested black macaques in primary forest on Bacan is best explained by the high carrying capacity found in primary forest. The lower food quantity and quality of food resources found in logged forest correlated with lower primate densities compared to primary forest. However, the large population of macaques in logged forest demonstrates the conservation value of such forest. Densities on Sulawesi at Tangkoko-Batuangas-DuaSudara Nature Reserve (TBDS) showed a continuing decline since earlier surveys. Primate densities were highest near the protected center of Tangkoko Reserve (66.7 individuals/km²). The peripheral areas of Batuangas and DuaSudara, even though adjacent and continuous, showed lower population densities of 46.4 and 23.5 individuals/km², respectively. The best explanation for the continued decline of Macaca nigra populations at TBDS is hunting. Unless conservation measures are implemented immediately, M. nigra on Sulawesi risks extinction in the near future. Am. J. Primatol. 44:89–106, 1998. © 1998 Wiley-Liss, Inc.     

Leaf gas exchange of trees in old-growth and young secondary forest stands in Sulawesi, Indonesia

In the tropics, old-growth forests are converted to other land cover types at a high rate and young secondary forest may gain in importance. Information on associated changes in leaf gas exchange and other leaf traits can be valuable for modelling biogeochemical fluxes under altered land-use patterns. We studied in situ photosynthetic parameters and stomatal conductance for water vapour in eight abundant tree species of young secondary forest and eight tree species of natural old-growth forest in Central Sulawesi, Indonesia. In sun leaves, the average maximal stomatal conductance (g _smax) in the secondary forest (SF) species was 2.1 times higher than in the old-growth forest (OGF) species. Species with a high g _smax reduced g _s sharply when vapour pressure deficit of the air increased, whereas species with a low g _smax were much less sensitive to air humidity. For area-based photosynthetic capacity (A _max-area), the SF species had a 2.3 times higher average than the OGF species. For both, g _smax and A _max-area the variation among species was higher in the OGF than in the SF. When all tree species (n=16) are considered, species means of specific leaf area (SLA), leaf N concentration and leaf P concentration were significantly correlated with g _smax and A _max-area. The strong correlation between A _max-area and foliar P (r ²=0.8) is remarkable as the alluvial soils in the study region are rich in nutrients. If the eight OGF species are analysed separately, the only significant correlation was observed between SLA and mass-based A _max; in the SF species strong correlations were found between leaf size and A _max-area and g _smax. These results show that the conversion of old-growth forest to young secondary forest in Sulawesi significantly alters tree leaf gas exchange characteristics and that chemical and structural leaf traits can be used for the prediction of these changes. The best correlations between leaf gas exchange parameters and leaf traits were obtained by different traits in the SF species, the OGF species and the entire pool of studied species.     

Ectoparasites of commensal rodents in Sulawesi Utara, Indonesia, with notes on species of medical importance

Ectoparasite records are presented for four species of commensal murid rodents (Rattus rattus palelae Miller & Hollister, R. argentiventer (Robinson & Kloss), R. exulans (Peale) and Mus musculus castaneus Waterhouse) in Sulawesi Utara, with particular reference to the potential for these arthropods to bite and transmit pathogens to humans. The flea, Xenopsylla cheopis (Rothschild), was most common on R.r. palelae and is capable of transmitting plague and other pathogens to humans although no current foci for these diseases are known in Sulawesi. Hoplopleura pacifica Ewing and Polyplax spinulosa (Burmeister) sucking lice parasitized all three Rattus species although H. pacifica was mainly associated with R. exulans and P. spinulosa with R.r. palelae. These lice do not bite humans but may be intramurid vectors of murine typhus and other zoonoses. The mites Laelaps echidnina Berlese and L. nuttalli Hirst were both collected; the latter was recorded from all four murid species, mainly R. exulans. The mite Ornithonyssus bacoti Hirst was rare. Only one chigger mite, Walchiella oudemansi (Walch), was retrieved from murids (from R. exulans) and a single Leptotrombidium deliense (Walch) chigger was taken from a human subject. Although L. deliense is a significant vector of scrub typhus, a disease known from Sulawesi, the L. deliense-R. argentiventer relationship frequently noted in the ecology of this rickettsial disease, was not evident in this survey. Other ectoparasites collected from murids were the ticks, Ixodes granulatus Supino (first record for Sulawesi), Haemaphysalis sp. and Dermacentor sp., the mites Myocoptes musculinus (Koch) and Listrophoroides cucullatus (Trouessart), acarids and a uropodid.(ABSTRACT TRUNCATED AT 250 WORDS)     

Dispersal and population genetic structure of Telmatherina antoniae,an endemic freshwater Sailfin silverside from Sulawesi,Indonesia

Population genetic structure in the presence of substantial dispersal provides a unique perspective on the evolution of reproductive isolation. We sampled Telmatherina antoniae, an endemic fish species, at 10 sites in Lake Matano, Indonesia. Significant genetic structure (F_ST = 0.03) was found, despite a migration rate of 10.2% and a mean dispersal distance of 13.6 km, estimated by genotype assignment. Neither dispersal distance nor direction differed from random expectations, indicative of no dispersal barrier in Lake Matano. However, Bayesian genotype cluster assignment identified a population structure consisting of four to six clusters that did not coincide with sample site distribution, but explained two to three times more genetic variance than sample site. The mechanism for continued isolation of those genetic clusters is unknown, but assortative mating and temporal isolation are obvious candidates. Our results resolve the apparent paradox of population genetic structure coupled with frequent dispersal, and highlight the importance of considering cryptic genetic structure.     

Colonial "Reformation" in the Highlands of Central Sulawesi, Indonesia, 1892-1995

Colonial "Reformation" in the Highlands of Central Sulawesi, Indonesia, 1892-1995. Albert Schrauwers. Toronto: University of Toronto Press, 2000. 279 pp.     

Tree diversity and phytogeographical patterns of tropical high mountain rain forests in Central Sulawesi,Indonesia

Tropical high mountain forests in Lore Lindu National Park, Sulawesi, Indonesia, were described by their floristic composition and the importance of tree families (Family importance values, FIV), based on tree inventories conducted on 4 plots (each 0.24 ha) in old-growth forest stands at c. 1800 and 2400 m a.s.l. (mid- and upper montane elevations). To identify general patterns and regional peculiarities of the forests in the SE Asian and SW Pacific context, the biogeography of the tree species was analysed using distribution records. Out of the total of 87 tree species, only 18 species were found at both elevational zones. The discovery of new species and new distribution records (28% of the data set) highlights the deficiencies in the taxonomic and distribution data for Sulawesi. Sulawesi endemism rate was 20%. In the mid-montane Fagaceae–Myrtaceae forests, Lithocarpus spp. (Fagaceae) were overall important (4 spp. occupying around half of the total basal area) and the Myrtaceae the most species rich (8 spp.), thus showing typical features of Malesian montane forests. The upper montane conifer-Myrtaceae forest contained several high mountain tree taxa and showed affinity to the forests of New Guinea. The mountain flora comprised both eastern and western Malesian elements, with the nearest neighbouring islands Borneo and Maluku both sharing species with Sulawesi, reflecting the complex palaeogeography of the island. A separate analysis showed the mid-montane forest to possess greatest biogeographical affinity to Borneo/western Malesia, and the upper montane forest had a number of typical elements of Papuasia/eastern Malesia and the Phillipines, which may be a result of historical patterns in land connection and the emergence of mountain ranges.     

Conservation Value of Cacao Agroforestry Systems for Terrestrial Herbaceous Species in Central Sulawesi,Indonesia

Tropical secondary forest and agroforestry systems have been identified as important refuges for the local species diversity of birds and other animal groups, but little is known about the importance of these systems for terrestrial herbs. In particular, few studies report how the conversion from tropical forest to technified cacao plantation affects the species richness and the community structure of herbs. We conducted surveys in 43 cacao plantations along the border of the Lore Lindu National Park in Central Sulawesi, ranging from agroforests to technified cacao, categorizing the plantations as rustic cacao, planted shade cacao, and technified cacao. We recorded 91 herb species. Of the 74 species determined to species level, 21 were also found in natural forests, while 53 were recorded only in agricultural habitats. Araceae was the most forest‐dependent plant family while Asteraceae included the highest number of nonforest species. Overall, the presence of forest species was confined to moderately intensively managed rustic and planted shaded plantations. Distance from the forest, which has been identified as a crucial parameter for the diversity and composition of other taxa in cacao agroforests, only played a minimal role for herbs. Our study suggests that native forest herbs maybe more vulnerable to forest conversion than animal groups. The intensification of cacao plantation management increases the presence of weedy species to the detriment of native forest species.     

Mimicry and foraging behaviour of two tropical sand-flat octopus species off North Sulawesi, Indonesia

The so-called 'mimic octopuses' of tropical Indonesia are reputed to mimic up to 13 species of other local marine organisms. We tested for mimicry by allowing individuals of two species of octopus to habituate to divers, then observing and filming two species continuously as they foraged daily in the same open, featureless volcanic sand habitat. Mimicry of a local, abundant flounder occurred commonly during 5 days of natural foraging: nearly 500 episodes were analysed. Both octopus species mimicked the shape, swimming actions, speed, duration, and sometimes the coloration of swimming flounders. During flounder mimicry, octopuses were actively moving and conspicuous, whereas immediately before and after flounder mimicry, they were camouflaged and motionless (sitting or very slowly crawling). Furthermore, when motionless, octopuses assumed body patterns and postures that resembled small sponges, tube-worm tubes, or colonial tunicates, which were among the few objects in the open sand habitat. The key finding was that octopuses used flounder mimicry only when their movement would give away camouflage in this open habitat. In all cases, octopuses used mimicry as a primary defense. Several interactions with fishes and stomatopods were filmed and typical secondary defense behaviours, not mimicry, were used by the octopuses. Foraging occurred twice per day and two tactile feeding tactics were used. Dens and food were not limiting; thus, we observed a highly unusual circumstance of a guild of small, long-armed octopus species that shared the same habitat, den sources, food, activity period, and some behaviours.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 23–38.     

Loss of habitats,naturalness and species diversity in Eurasian forest landscapes

Man has exploited land and forests in Western and Central Europe longer and more intensively than in Northern Europe and further east in Eurasia. We estimated forest naturalness and modelled expected biodiversity loss in seven different landscapes (2500 km² each) in the Netherlands, Sweden, Finland, Poland, St. Petersburg (Western European Russia), Perm (Eastern European Russia), and Irkutsk (Central Siberia) across the distribution of Pinus sylvestris L. in Eurasia. Field inventories showed that the mean living tree volumes were relatively similar in the studied sites, but the volumes of dead wood differed greatly. In Irkutsk and Perm the volume of dead trees per ha was about 5–10 times larger than in Central and Western European regions. The studied forests were generally young in all regions except for Irkutsk, where about half of the study plots had trees older than 120 years. Signs of recent forest fires were found almost exclusively on Russian sites. According to Landsat satellite image-based land-cover classifications the amount of remaining forest habitat in the studied landscapes varied from 25% in the Netherlands to 93% in Irkutsk. Estimated by forest patch size and density of cut stumps, forests were also more fragmented and heavily managed in the western study landscapes compared to eastern ones. Based on species–area relationship functions, we calculated that the proportion of forest-dwelling species already extinct or expected to become extinct due to habitat loss ranges from 1–2% in Irkutsk to 13–24% in the Netherlands study landscape. For saproxylic species, which depend on dead wood, the extinction estimates were calculated based on remaining dead wood volume in the landscape. The modelled expected loss of saproxylic species ranged from 7–14% in Irkutsk to 35–58% in the Netherlands.     

Dissimilar patterns of genetic variation in two insular endemic plants sharing species characteristics,distribution, habitat,and ecological history

Camssonia guadalupensis ssp. clementina (Onagraceae) and Cryptantha traskiae (Boraginaceae) share many species characteristics that affect levels and pattern of genetic variation. In addition, they have a similar distribution, are found in the same habitat, and share ecological history by virtue of co-occurring at nearly all of the locations at which they are found. All populations of the two taxa on San Clemente Island were surveyed for genotypes at 16 allozyme loci, revealing generally low levels of genetic variation. However, Cam. g. clementina has more genetic variation than Cryp. traskiae at the taxon level (P = 37.5 versus 18.8, A = 1.69 versus 1.31, and H_E = 0.088 versus 0.003) and significantly more alleles and higher heterozygosity within populations (A = 1.09 versus 1.03, A_E = 1.02 versus 1.00; H_O = 0.013 versus 0.001 and H_E = 0.017 versus 0.003). Some populations of each taxon are monomorphic at all loci. Much greater differentiation among populations occurs in Cam. g. clementina (G_ST = 0.810) than in Cryp. traskiae (G_ST = 0.042). Locations of the more variable populations of each taxon do not coincide, and the pattern of genetic similarity of localities differs in the two taxa. Striking population genetic differences exist between these taxa despite shared species characteristics, distribution, habitat, and recent history.