Biogeography of the Indonesian archipelago: mitochondrial DNA variation in the fruit bat, Eonycteris spelaea

The fruit bat, Eonycteris spelaea , occurs from India through the Philippines to the southeast limit of its distribution in the Lesser Sunda islands of Indonesia. Mitochondrial DNA (mtDNA) variation was examined in Indonesian E. spelaea island populations by amplification of the D-loop and digestion with restriction endonucleases. In addition, microgeographic variation was assessed by investigation of three cave populations within one island. A total of 24 genotypes, comprising two broad clades, was detected. The pattern of mtDNA variation reflects the colonization history of E. spelaea with estimates of haplotype and sequence diversity highest in the older western populations and lowest at the eastern periphery of the species' distribution. These findings may also be associated with an environmental cline from west to east. There is also evidence that genetic distance between populations reflects geographic relationships, especially historical connectedness, as measured by Pleistocene sea-crossing distances. At the microgeographic level, cave populations were heterogeneous and composed of diverse lineages suggesting restricted local interchange.     

Characterization of microsatellite loci in the lesser dawn bat (Eonycteris spelaea)

We constructed a genomic DNA library enriched for CA repeat motifs in Eonycteris spelaea. Nine microsatellite loci were isolated and tested on a population of 39 samples from Yunnan Province, China. These nine loci had three to 22 alleles per locus. Observed and expected heterozygosity values ranged from 0.079 to 0.963 and from 0.078 to 0.959. Two loci revealed significant departure from Hardy-Weinberg equilibrium and no linkage disequilibrium was found between loci pairs. These microsatellites can be a powerful molecular tool for population-level studies of E. spelaea.     

Allozyme variation in turbot (Psetta maxima) and brill (Scophthalmus rhombus) (Osteichthyes, Pleuronectoformes, Scophthalmidae) throughout their range in Europe

Two species of coastal flatfish (brill and turbot, Scophthalmidae) were analysed electrophoretically at 17 common enzymatic loci in samples taken from 11 sites representing the species ranges in Europe. Brill showed a mean heterozygosity ( H ) of 0·11 while that of turbot was 0·02. The virtual absence of genetic diversity in turbot is probably due to a very low evolutionary rate, and provides little evidence for population substructure even if various signs indicate the possibility of a hidden differentiation (presence of the taxon maoetica in the Black Sea and differentiation of a species-specific cestode parasite on either side of the Strait of Gibraltar). On the other hand, the weak geographic structure in brill seems to result from rapid recolonization following the last ice age.     

A phylogeny for grasshoppers of the genus Chitaura (Orthoptera: Acrididae) from Sulawesi, Indonesia, based on mitochondrial DNA sequence data

The Indonesian island of Sulawesi occupies a central position within the biogeographically complex region known as Wallacea. Its fauna is characterised by a high rate of endemism and a patchwork distribution of taxa within the island. The grasshopper genus Chitaura is a good example having at least ten endemic species with predominantly parapatric distributions. It can be used as a model for determining the origins of Sulawesi taxa and the within-island evolution that has led to the present patterns of distribution. Here we present a phylogenetic hypothesis for 28 individuals within the genus, including individuals of one species from Java and two from the Moluccas, based on DNA sequences of the mitochondrial cytochrome oxidase 1 gene. Frequent sequence heteroplasmy was observed. The phylogenetic hypothesis is consistent with recent interpretations of the geological history of Sulawesi suggesting separate evolution on the island for 7–14 Myr, possibly since South Sulawesi was connected to Borneo. Within the island, the pattern of genetic divergence is dominated by a strong correlation with geographic distance, with exceptions indicating past or present barriers to dispersal. Colonisation of the Moluccas from North or Central Sulawesi is implied. Levels of genetic divergence are compared with distribution patterns of colour morphs and with possible effects of tectonic movements in the Cenozoic, or Pleistocene climatic, vegetational and sea-level changes.     

Flight is the key to postprandial blood glucose balance in the fruit bats Eonycteris spelaea and Cynopterus sphinx

Excessive sugar consumption could lead to high blood glucose levels that are harmful to mammalian health and life. Despite consuming large amounts of sugar‐rich food, fruit bats have a longer lifespan, raising the question of how these bats overcome potential hyperglycemia. We investigated the change of blood glucose level in nectar‐feeding bats (Eonycteris spelaea) and fruit‐eating bats (Cynopterus sphinx) via adjusting their sugar intake and time of flight. We found that the maximum blood glucose level of C. sphinx was higher than 24 mmol/L that is considered to be pathological in other mammals. After C. sphinx bats spent approximately 75% of their time to fly, their blood glucose levels dropped markedly, and the blood glucose of E. spelaea fell to the fast levels after they spent 70% time of fly. Thus, the level of blood glucose elevated with the quantity of sugar intake but declined with the time of flight. Our results indicate that high‐intensive flight is a key regulator for blood glucose homeostasis during foraging. High‐intensive flight may confer benefits to the fruit bats in foraging success and behavioral interactions and increases the efficiency of pollen and seed disposal mediated by bats.     

Allozyme Variation within and between Populations of Coreopsis intermedia (Asteraceae)

Abstract Enzyme electrophoresis was used to measure genetic diversity in two populations of Coreopsis intermedia of section Coreopsis , a rare species endemic to a small area of Texas and Louisiana. Genetic diversity is higher in C. intermedia than has been reported for many other geographically restricted species. C. intermedia contains levels of genetic variation comparable to those of other species in section Coreopsis , and a greater amount of diversity than either the rare congener, C. latifolia (section Silphidium ), or the Florida endemic, C. leavenworthii (section Calliopsis ). The two populations, although greatly different in size, exhibited similar levels of genetic diversity.     

Gut morphology and morphometry in the epauletted Wahlberg's fruit bat (Epomophorus wahlbergi, Sundevall, 1846)

The morphological adaptations of the fruit bat small intestine to which the high functional efficiency could be related and the possible landmarks delineating the various parts of the gut were examined. The stomach was the carnivorous type with large rugae spanning the entire luminal aspect down to the pyloric sphincter, which was reflected internally as a prominent fold. Externally, the intestine was a continuous tube uninterrupted by any structures. The cranial fifth of the small gut had long, branching and anastomosing villi, which caudally turned to finger-like discrete structures that became rather short and stumpy and diminished at the beginning of the colon. The colon had longitudinal folds that were macroscopically discernible from the mucosal aspect of the opened intestine and that continued into the rectum. The small gut formed 94% of the whole intestinal length, the colon and the rectum taking 4 and 2%, respectively. Ultrastructurally, the enterocyte showed a prominent brush border and the lateral membranes were modified into numerous tortuous interdigitating processes. Adjacent enterocytes were joined by these processes through desmosomes. The processes also participated in pinocytotic fluid uptake from the intercellular spaces with resultant numerous intracellular vacuoles of varied sizes. Solutes absorbed into the cells were probably first passed into the intercellular compartment to create a concentration gradient thus enhancing further absorption into the cell. We conclude that the uniquely elaborate ultrastructure of the enteric epithelium coupled with the vast microvillous surface areas reported elsewhere are partly responsible for the very high absorption rates reported in the fruit bat small intestine.     

Genetic structure and environmental heterogeneity in the European hake (Merluccius merluccius)

This study aimed at assessing the genetic structure and the state of the stocks of the European hake (Merluccius merluccius). To this end, 15 samples were taken from the whole range of the species and analysed using allozymes. Since 11 samples were taken from the poorly studied Mediterranean Sea, the results obtained provided a complete picture of the hake's genetic structure and an initial insight into its relationships with environmental features. Atlantic and Mediterranean hake populations are separated by the Almeria-Oran front. This area has been proved to be the boundary between Atlantic and Mediterranean stocks of many marine organisms, but some doubt exists concerning the efficaciousness of the local gyres as barriers to the gene flow. Our data have evidenced a latitudinal cline at loci Gapdh and Gpi-2 within the Mediterranean Sea, with a further steep change across the Almeria-Oran front. The genetic pattern showed a strong correlation with the values of the salinity both at the surface and at -320 m and of the salinity + temperature at the surface, suggesting a role for these parameters in maintaining the genetic differentiation among the two population groups through selective processes. Finally, the levels of genetic variability were found to be slightly lower in the depleted Atlantic stock than in the Mediterranean one.     

Population genetics and the conservation status of the threatened Iberian steppe grass Puccinellia pungens (Pau) Paunero (Poaceae)

Puccinellia pungens (Pau) Paunero is a narrowly endemic grass found in two continental saline lagoons of north-eastern Spain. This rare plant has been classified as 'at risk of extinction' in several national and European catalogues of endangered species. Recent demographic studies indicate that population sizes greatly exceed several million individuals, challenging that threat category. Our genetic analysis, based on allozymes, has shown that in spite of the large population sizes, very low levels of genetic variation were found in P. pungens . Genetic variation was similar in most populations, but the largest, Gallocanta lagoon as a whole, had less variation (35% polymorphic loci, 1.4 alleles/locus, H _T = 0.038) than the more restricted Royuela range (45% polymorphic loci, 1.5 alleles/locus, H _T = 0.056), suggesting a recent population expansion of the Gallocanta populations from few founder lines. The low genetic distances among populations also suggest a recent divergence. The low genetic variation observed cannot be explained fully by eventual clonal spread and rare seedling establishment in the hypersaline environment. This low variation seems to result from extreme recent population bottlenecks as a consequence of habitat conversion to agricultural fields. In the light of our data, it seems unlikely that reinforcement of populations could increase the genetic diversity of the populations. Hence, conservation efforts should focus on avoiding further habitat loss of this endangered steppe grass species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 269–281.     

Conservation genetics of peripheral populations of the mygalomorph spider Atypus affinis (Atypidae) in northern Europe

Genetic diversity and differentiation were studied in peripheral populations of the recently rediscovered purse-web spider Atypus affinis in Denmark and Sweden using allozyme electrophoresis. Because of the very narrow environmental niche exploited by this species, only a limited number of potential habitats are available. Furthermore, fragmentation has reduced the number and size of potential habitats within the last century. The level of genetic diversity in A. affinis was intermediate compared with other nonsocial spiders, but low compared with invertebrates in general. Significant genetic differentiation was found within distances of only 1-10 km with FST estimates ranging from 0.020 to 0.075. Within distances of 30-60 km FST ranged from 0.081 to 0.312. Hierarchical FST revealed that genetic variability was partitioned at 89.8% within populations, 9.5% among populations within regions and only 0.7% among the four main regions in Denmark and Sweden. Comparing the result of the genetic analysis with the life history of A. affinis, it is concluded that the level of successful dispersal is low, and that the species has not recently reinvaded northern Europe but prevailed undiscovered for several decades. Finally, it is suggested that the genetic scenario found for A. affinis might represent that for a wide range of other arthropods with similar life history characteristics.     

Multiple levels of allopatric divergence in the endemic Philippine fruit bat Haplonycteris fischeri (Pteropodidae)

As part of a larger comparative phylogeographical study of Philippine fruit bats, I used fragments of the mitochondrial genes cytochrome  b and ND2 to investigate phylogeography and diversification in Haplonycteris fischeri , a pteropodid bat endemic to the Philippines but widespread within the archipelago. Genetic diversity in H. fischeri was extremely high in these commonly studied genes, with 101 unique haplotypes in 123 sequenced individuals, although small, continuously isolated islands had less diversity than had large island complexes. Seven monophyletic groups and one paraphyletic group were restricted to individual islands, groups of islands, or parts of islands. Each Pleistocene island complex had a single resident monophyletic lineage; these five groups were separated by approximately 6–8% sequence divergence and apparently have been diverging for 4–6 Myr. Within island groups, monophyletic lineages on some individual islands suggest that current ocean channels have also been barriers to gene flow; in some cases, multiple allopatric clades were present on single islands. Basal divergence dates were estimated to be in the early Pliocene, and most diversification was apparently connected to the ongoing geological evolution of the Philippines. Geological history and current geography interact with ecology to cause substantial genetic differentiation within this primary forest-specialist species.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 329–349.     

Revision of the coral genus Acropora (Scleractinia: Astrocoeniina: Acroporidae) in Indonesia

The coral genus Acropora is reviewed from Indonesia for the first time, following detailed collections made at 131 sites and additional material collected from approximately 40 sites throughout the archipelago during the period 1993–6. Eighty-three species are recorded, four of these ( Acropora halmaherae, A. awi, A. plumosa and A. simplex ) new to science, six first described in 1994 and six in 1997. Records are compared with specimen-based records from localities worldwide. The species of Acrokora occurring in Indonesian waters include five recorded only from the Indian Ocean and Indonesia, seven recorded only from the Pacific Ocean, South China Sea and Indonesia, and a further 10 species apparently endemic to Indonesia, as well as widespread Indo-Pacific species. Two species ( A. jacquelineae Wallace, 1994 and A. batunai Wallace, 1996) are recorded only from north central Indonesia and Papua New Guinea, and two species ( A. russelli Wallace, 1994 and A. turaki Wallace, 1994) only from north central Indonesia and north western Australia. The findings contribute to a new view of the corals of the Indo-Pacific 'centre of diversity' as a composite fauna with origins in a number of events in space and time.     

Cryptic speciation, genetic diversity and gene flow in the California turret spider Atypoides riversi (Araneae: Antrodiaetidae)

Populations of the turret spider Atypoides riversi from eight central Californian sites were compared based on variation at ten allozyme loci. Multidimensional scaling of interpopulation genetic distances defined four population units (Coast Range, Sierran, Valley, Jenness Camp), corroborating the distinctness of Coast Range and Sierran populations indicated by a prior study. While the species status of these units has yet to be determined, Jenness Camp is the most likely to represent a new species, given its clear genetic uniqueness (two fixed allelic differences). Populations in all units were generally in Hardy–Weinberg equilibrium with no evidence of inbreeding, though variability was minimal (mean H _o = 2.8%, mean P  = 15.4%). Reduced variability in these populations may be the result of repeated bottlenecks, environmental homogeneity, and/or directional selection. Interpopulation differentiation within units was significant in the absence of intervening forest habitat and was substantially less in its presence, indicating that gene flow is likely only when forest corridors exist. To foster preservation of the existing gene pool and enhance participation in it, management of the units of At. riversi should focus on maintaining as many populations in situ as possible and facilitating connections between them, while also creating or restoring habitat for potential colonization.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 27–37.     

Genetic variation in the Desert Springsnail (Tryonia porrecta): implications for reproductive mode and dispersal

Allozymes and mitochondrial cytochrome c oxidase subunit I (mtCOI) sequences were analysed to determine whether populations of the western North American gastropod Tryonia porrecta (from California, Nevada, Utah, and northwest Mexico) are strongly differentiated in accordance with traditional interpretation of regional fauna as ancient relicts inhabiting isolated fragments of late Tertiary palaeodrainages. These data were also used to assess whether this species, for which males have not been recorded, is a rare example of a molluscan parthenogen. Both data sets strongly supported monophyly of T. porrecta populations. Five of the nine sampled populations consisted of a single monoallelic allozyme genotype while the others contained two to 10 distinct genotypes. Allozymic data for genetically diverse Utah populations provided evidence of clonal and sexual reproduction. mtCOI haplotypes of T. porrecta formed two subgroups which differed by 1.99-2.60%. The common haplotype was found in seven populations with rare haplotypes observed in single populations. Based on these results and an available mtCOI molecular clock for related hydrobiid snails, T. porrecta is interpreted as a primarily parthenogenetic species that undergoes occasional sexual reproduction and has accumulated substantial diversity following its mid-Pliocene to mid-Pleistocene origin. Our results also suggest that the distribution of present-day populations of these gill-breathing snails did not result from fragmentation of an ancient, well-integrated drainage but instead reflects overland colonization of habitats which only recently became available following desiccation of late Quaternary pluvial lakes.     

Responses to light in two blind cave fishes (Amblyopsis spelaea and Typhlichthys subterraneus) (Pisces: Amblyopsidae)

Explanations for the phenotypical features resulting from colonization of subterranean environments have always been a source of controversy. Although a great number of cave organisms are blind, they nevertheless display responses to light. The interpretation of this phototactic responsiveness in cave-dwelling animals may provide clues on the general issue of evolution of behavior in parallel with specialized structures. We studied the phototactic responses in two amblyopsid fishes of North America and found responses to light only in the species reported to have a functional pineal organ. Our findings are consistent with the hypotheses that (1) adaptation to the cave environment is a gradual process and (2) responsiveness to light in cave fishes may best be understood as a relict character, one that exists in an environment where it may never be expressed.     

Geographical variation in the morphology of four snake species from the Lesser Sunda Islands, eastern Indonesia

The morphology of three colubrid snakes, Elaphe subradiata, Psammodynastes pulverulentus, Lycodon aulicus capucinus , and the viper Trimeresurus albolabris , were examined from populations throughout the Lesser Sunda islands using canonical variate analysis. The Lesser Sundas form the western part of two extensive Island chains, the Banda Arcs, that lie between the large islands on the Sunda (Asian) and Sahul (Australian) continental shelves. The snakes of the Lesser Sunda islands show considerable intraspecific variation in morphology. Populations of Trimeresurus albolabris from Wetar and Psammodynastes pulverulentus from Alor show the greatest morphological separation from conspecifics. The most pronounced morphological differentiation in multidimensional space occurs between populations on islands that remained separated throughout the Pleistocene, when sea levels were about 120 m lower than present. Consensus tree evaluation of Mahalanobis distance for populations of these four species and Dendrelaphis pubis on the islands of Lombok, Sumba, Flores, Lembata and Alor, supports the finding that populations from islands that remained isolated throughout the Pleistocene by sea barriers show the greatest morphological divergence.     

Wing morphology and flight development in the short-nosed fruit bat Cynopterus sphinx

Postnatal changes in wing morphology, flight development and aerodynamics were studied in captive free-flying short-nosed fruit bats, Cynopterus sphinx. Pups were reluctant to move until 25 days of age and started fluttering at the mean age of 40 days. The wingspan and wing area increased linearly until 45 days of age by which time the young bats exhibited clumsy flight with gentle turns. At birth, C. sphinx had less-developed handwings compared to armwings; however, the handwing developed faster than the armwing during the postnatal period. Young bats achieved sustained flight at 55 days of age. Wing loading decreased linearly until 35 days of age and thereafter increased to a maximum of 12.82 Nm(-2) at 125 days of age. The logistic equation fitted the postnatal changes in wingspan and wing area better than the Gompertz and von Bertalanffy equations. The predicted minimum power speed (V(mp)) and maximum range speed (V(mr)) decreased until the onset of flight and thereafter the V(mp) and V(mr) increased linearly and approached 96.2% and 96.4%, respectively, of the speed of postpartum females at the age of 125 days. The requirement of minimum flight power (P(mp)) and maximum range power (P(mr)) increased until 85 days of age and thereafter stabilised. The minimum theoretical radius of banked turn (r(min)) decreased until 35 days of age and thereafter increased linearly and attained 86.5% of the r(min) of postpartum females at the age of 125 days.     

Functional morphology, ecology, and evolution of the Scaphitaceae Gill, 1871 (cephalopoda)

Scaphitids are heteromorph ammonites exhibiting morphologicaltrends counter to of those of the other main heteromorph ammonitegroups. These trends include the shortening of the body chamber,lateral compression of the whorl, closure of the coil, and moreregular, spiral coiling. Scaphitid-like morphologies may haveappeared in other heteromorphs, but the Scaphitaceae are monophyletic.The most primitive scaphitids are known from the Albian, butan important radiation occurred in the Western Interior Seawayof North America from the Santonian to Maastrichtian, some ofwhich spread to the Old World. The scaphitid morphology is consistentwith improved swimming abilities, but scaphitids remained associatedwith the seafloor and are best considered to have been nektobenthossimilar to modern nautiluses.     

Understanding population history for conservation purposes: population genetics of Saxifraga aizoides (Saxifragaceae) in the lowlands and lower mountains north of the Alps

Several alpine species have outlying populations in the lowlands and lower mountains north of the Alps. These small, isolated populations are usually described as either (1) glacial relics, (2) descendants from populations living on forelands and moraines during the ice ages, or (3) populations founded by long-distance dispersal after glaciation. A floristic survey of the historic and present distributions and an allozyme investigation were performed on one of these relic species, Saxifraga aizoides. The species was historically more abundant and had more stations in more regions of northeastern Switzerland. The former population structures within regions, nowadays destroyed, were still reflected in distinct and high regional genetic diversity and variation. There was weak evidence of increased inbreeding in outlying populations, but populations did not deviate from Hardy-Weinberg equilibrium. No geographic pattern of genetic variation above the regional scale (>10 km) was found. Based on the spatial and genetic structures found, it was not possible to discriminate between the abovementioned hypotheses. Nevertheless, the study shows how a thorough evaluation of distribution and abundance data aids the interpretation of genetic data with respect to population history, biogeography, and conservation biology.