A molecular phylogenetic analysis of the family Rhacophoridae with an emphasis on the Asian and African genera

Using characters from mitochondrial DNA to construct maximum parsimony and maximum likelihood trees, we performed a phylogenetic analysis on representative species of 14 genera: 12 that belong to the treefrog family Rhacophoridae and two, Amolops and Rana, that are not rhacophorids. Our results support a phylogenetic hypothesis that depicts a monophyletic family Rhacophoridae. In this family, the Malagasy genera Aglyptodactylus, Boophis, Mantella, and Mantidactylus form a well-supported sister clade to all other rhacophorid genera, and Mantella is the sister taxon to Mantidactylus. Within the Asian/African genera, the genus Buergeria forms a well-supported clade of four species. The genera, except for Chirixalus, are generally monophyletic. An exception to this is that Polypedates dennysii clusters with species of Rhacophorus, suggesting that the taxonomy of the rhacophorids should be revised to reflect this relationship. Chirixalus is not monophyletic. Unexpectedly, there is strong support for Chirixalus doriae from Southeast Asia forming a clade with species of the African genus Chiromantis, suggesting that Chiromantis dispersed to Africa from Asia. Also, there is strong support for the sister taxon relationship of Chirixalus eiffingeri and Chirixalus idiootocus apart from other congeners.     

侧条跳树蛙的分布新纪录及其分布区域分析

2008年5月30日,在广东黑石顶自然保护区采集到5只侧条跳树蛙(Chirixalus vittatus),为广东省两栖动物新纪录.通过对侧条跳树蛙分布区域分析,认为十万大山至云开大山之间的山区是侧条跳树蛙在中国大陆的主要分布区,侧条跳树蛙的现代分布属中印半岛-中国南部热湿型.     

The thermal environment of arboreal pools and its effects on the metabolism of the arboreal, oophagous tadpoles of a Taiwanese tree frog, Chirixalus eiffingeri (Anura: Rhacophoridae).

We have studied seasonal and diurnal fluctuations of water temperature in bamboo stumps and the effect of temperature on the energy metabolism of arboreal, oophagous tadpoles of Chirixalus eiffingeri. We collected tadpoles (Gosner stage 28-29) in February and August from Chitou, Taiwan and acclimated them to 12 and 22 degrees C. Using a closed system, we measured tadpole oxygen consumption (V.O(2)) at 12, 17 and 22 degrees C. The water temperature was lowest in February (11-13 degrees C), increased rapidly during March and April and was highest from May to August (20-24 degrees C). Diel fluctuations in the temperature of the pools of water in bamboo stumps mirrored fluctuations in air temperature. Tadpoles collected in February and August exhibited metabolic compensation in that tadpoles acclimated at 12 degrees C had significantly higher V.O(2) than those acclimated at 22 degrees C. There are at least two possible explanations for the presence of metabolic compensation in C. eiffingeri tadpoles. Firstly, the larval period of C. eiffingeri ranges from 40 to 78 days, a tadpole could experience relatively large fluctuations in body temperature (up to 10 degrees C) during the development. As a result, C. eiffingeri tadpoles most likely evolved metabolic compensation to maintain activity levels under different thermal environments. Secondly, because arboreal pools are small, thermally unstratified, aquatic microhabitats, tadpoles are unable to behaviorally select preferred temperatures. As a result, metabolic compensation allows tadpoles to regulate their physiological functions.     

Isolation and characterization of polymorphic microsatellite loci for parentage analysis in a rhacophorid tree frog (Chirixalus eiffingeri) with unusual parental care

Chirixalus eiffingeri is an arboreal breeding rhacophorid frog with unique parental care behaviours. We developed 11 polymorphic microsatellites as genetic makers for parentage analysis to resolve the ecology of parental care. The numbers of alleles per locus ranged from two to 17. The observed and expected heterozygosities averaged 0.433 and 0.656, respectively. Total exclusionary probability of these loci is 0.984 when no parental genotypes are known, and is 0.999 when one of the parental genotypes is known. The results indicate that the 11 markers should provide sufficient resolution for inferring genetic parentage in C. eiffingeri.     

The fatty acid composition of oophagous tadpoles (Chirixalus eiffingeri) fed conspecific or chicken egg yolk

We compared the lipid content and fatty acid composition of (1) the egg yolk of three anuran species (Chirixalus eiffingeri, Rhacophorus moltrechti and Buergeria robustus) and chicken eggs; and (2) C. eiffingeri tadpoles fed conspecific eggs or chicken egg yolk. Anuran and chicken egg yolk contained more non-polar than polar lipids but the proportions varied among species. Chicken egg yolk contained low amounts of 22:5n-3 in the polar lipid fraction, and B. robustus eggs did not contain any n-3 or n-6 non-polar lipids. The specific variation of lipid contents and fatty acid composition may relate to the maternal diet and/or breeding biology. In C. eiffingeri tadpoles that fed chicken yolk or frog egg yolk, the dominant components of polar and non-polar lipids were 16:0, 18:0, 18:1, and 18:2n-6, or 20:4n-6 fatty acids. C. eiffingeri eggs contained more n-3 fatty acids (e.g. 18:3n-3 and 20:5n-3) than chicken egg yolk, and tadpoles fed conspecific eggs contained more of these fatty acids than tadpoles fed chicken egg yolk. The compositional differences in the fatty acids between C. eiffingeri tadpoles that fed frog egg or chicken egg yolk are the reflection of the variation in the dietary sources. Our results suggest a direct incorporation of fatty acids into the body without or minimal modification, which provide an important insight into the physiological aspects of cannibalism.     

The ecology of male egg attendance in an arboreal breeding frog, Chirixalus eiffingeri (Anura: Rhacophoridae), from Taiwan

Field observations were conducted on egg attendance in Chirixalus eiffingeri from April to August 2003. Parental attendance during embryonic development was performed exclusively by males. The frequency of egg attendance was low (27%), but it had a distinct diel pattern in which males were observed to attend eggs more frequently at night than during the day. Attendance frequency significantly decreased with increasing developmental stage of the embryos, but it was not statistically significantly related to clutch size. Field observations confirmed that male frogs actively moisten egg clutches using their ventral surfaces, presumably to prevent desiccation of egg clutches. The non-significant relationship between hatching success and frequency of egg attendance suggests that embryonic survival of C. eiffingeri is more than a function of egg attendance, and ecological and environmental factors, such as climate and characteristics of microhabitats, may also influence the survivorship of the embryos.     

Alternative migration and host parasitism strategies and their long-term stability in river lampreys from the River Endrick, Scotland

The stability of a discrete body size dimorphism of sexually mature river lamprey Lampetra fluviatilis from the River Endrick, Scotland, was examined over a 21 year period. Stable isotope analysis was used to test the hypothesis that the two size forms comprise individuals with differing migration and parasitic foraging strategies. Maturing river lamprey and the brook lamprey Lampetra planeri were trapped over 3 months each year in the periods 1983–1984 and 2004–2005. Brook lamprey catches and catches of both species combined showed no significant trend in catch rate with time. The catch rate of small body size river lamprey declined between 1983–1984 and 2004–2005 (although the difference did not reach statistical significance; P = 0·055). In contrast, there was a significant increase in the catch rate of the large body size river lamprey and as a consequence, a significant change in the relative proportion of each of the two river lamprey morphs over the study period. Analysis of the stable isotopes of C and N in muscle tissue showed that brook lamprey tissue derived its carbon from a freshwater source and had a δ¹³C more consistent with that of the River Endrick than with Loch Lomond. δ¹⁵N values for this species showed it to be feeding at the base of the food chain, consistent with filter feeding as an ammocoete. The large body size and the small body size river lamprey adults differed substantially in their δ¹³C values, with the small body size δ¹³C signature indicative of a freshwater carbon source and the large body size morph of a marine source. The small body size morph had a δ¹³C signature that was consistent with that of Loch Lomond powan Coregonus lavaretus suggesting that they share a common carbon source. The large body size morph was clearly feeding at a higher trophic level than the small body size morph. A single small body size river lamprey individual with typical morphology for that group, however, had C and N signatures that clustered with those of the large body size morphs. This individual had either migrated to sea to forage, as is typical for the species, or had been feeding on an anadromous fish with a strong marine C signature in fresh water. It is concluded that the body size dimorphism is indicative of a differential migration and foraging strategy in the parasitic phase of the life cycle of river lamprey at this site.     

Climatic effects and impacts of lakeshore bank designs on the activity of Chirixalus idiootocus in Yilan,Taiwan

Ecological engineering is the design of sustainable systems, consistent with ecological principles, which integrate human society with its natural environment for the benefit of both. In 2001, the government of Taiwan began to use ecological engineering as a therapy to nurse the damaged environment and found it worthwhile for further promotion for Taiwan's ecological environment. In recent years, the threat to amphibian animals is becoming more and more serious. The problem that concrete is mostly applied as a construction material on lakeshores in Taiwan has resulted in the obvious decrease of the habitats on which amphibian animals rely; hence, the need to develop the relevant design specification considering protecting the amphibian is imperative. Chirixalus idiootocus, an endemic species in Taiwan, is a small brown tree frog widely distributed in altitude ranging under 1500 m [Chen, W.S., 2003. 31 Frogs in Taiwan. Wild Bird Society of Taipei, pp. 62–63 (in Chinese)] with its unique breeding habitat [Kuramoto, M., Wang, C.S., 1987. A new Rhacophorid Treefrog from Taiwan, with comparisons to Chirixalus eiffingeri (Anura, Rhacophoridae). Copeia 4, 931–942]. Since its uniqueness and importance, it has been a good indicator to reflect the environmental change. However, the understanding of this species is still too little and the methods to protect them have not been developed yet. The goal of this study was to understand the impacts of different substrates, gradients, and humidities of lakebank slopes on the activity of Chirixalus idiootocus. Experiments were carried out on the samplings collected in the field at Chang-Pi Lake. Among the experimental results obtained in this study, the ability of jump height and jump length were 12.93–14.03 cm and 38.15–39.19 cm, respectively. The male group (♂) showed the best adhesive ability on awn grass (5.11 × 10⁻² N/g) at 45° slope, and the (♀) group performed the worst on sand (0.43 × 10⁻² N/g) at 75° slope. Female frogs showed significant worse performance than male in jump height, jump length, and adhesive abilities, so were considered a target reference to design bank slopes based on their vulnerable behavioral capacity. With better comprehensive results obtained in this study, suggestions for the bank slope designs were thus proposed as a useful reference for ecological engineering promotion in Taiwan.     

Complex selection on life-history traits and the maintenance of variation in exaggerated rostrum length in acorn weevils

Trophic interactions can trigger the development of exaggerated specialized characters and promote morphological diversification. For example, acorn weevils (genus Curculio) present strikingly long rostrums, which are used by females to perforate oviposition holes through the seed coat. Species exhibiting longer rostrums are known to exploit larger acorns, and therefore rostrum length is thought to be subject to selection to match the preferred acorn type. However, rostrum length is strongly correlated with body size, and morphological divergence could result from either selection on rostrum length for optimal food exploitation or from other pressures acting on body size. We collected infested acorns at oak forests where the large Curculio elephas and the small-bodied Curculio glandium co-occur. There were no interspecific differences in adult female body size to rostrum length allometric relationships, and rostrum length is equally correlated with body size in either species. MtDNA-based species identification showed that C. glandium larvae were present within acorns of all sizes, whereas C. elephas larvae were restricted to acorns above a minimum size, irrespective of oak species. Hence, exploitation of large acorns can hardly have triggered rostrum enlargement, as the small sized C. glandium adults (with short rostrums) could perforate and oviposit in both small and large acorns. Rather, increased rostrum length is probably a by-product of the larger body sizes of individuals emerging from bigger acorns, which allow increased larval size and enhance larval survival likelihood. Summarizing, when exaggerated feeding traits co-vary with other body features, interspecific morphological variability may result from contrasting selective pressures acting on these correlated characters.     

Day‐flying moths are smaller: evidence for ecological costs of being large

Research on evolutionary forces determining optimal body sizes has primarily relied on experimental evaluation of respective selective pressures. Accounting for among‐species variation through application of phylogenetic comparative methods is a complementary although little used approach. It enables the direct association of body size values with particular environments. Using phylogenetically explicit comparative analyses, we show that small body size is associated with diurnal (rather than nocturnal) activity of adults among temperate species of the moth family Geometridae. The association of an exclusively adult trait with species‐specific body size suggests that optimal body sizes are at least partly determined by the costs being a large adult, as opposed to the more frequently considered costs of attaining large size. It appears likely that size‐selective predation by insectivorous birds is the primary factor responsible for selection against large body size in day‐flying moths.     

Phylogenetic relationships among Rhacophorinae (Rhacophoridae, Anura, Amphibia), with an emphasis on the Chinese species

The partial nucleotide sequence of mitochondrial 12S and 16S rRNA genes was determined for 23 Chinese species of Rhacophoridae (Amphibia: Anura), representing four of the eight recognized genera. Using Buergeriinae as the outgroup, phylogenetic analyses (maximum parsimony, maximum likelihood, and Bayesian inference) were performed in combination with already published mitochondrial 12S and 16S sequences of Rhacophorinae frogs. In all cases, Philautus romeri Smith, 1953 is recovered as the sister taxon to all other Rhacophorinae, although the support values are weak. Chirixalus doriae Boulenger, 1893 is closer to Chiromantis [ Chiromantis rufescens (Günther, 1868) and Chiromantis xerampelina Peters, 1854] than to Chirixalus vittatus (Boulenger, 1887). The clade { Philautus odontotarsus Ye & Fei, 1993, [ Philautus idiootocus (Kuramoto & Wang, 1987), Kurixalus eiffingeri (Boettger, 1895)]} is recovered with strong support. The monophyly of Theloderma and Rhacophorus rhodopus Liu & Hu, 1959 is not supported. It is suggested that Philautus albopunctatus Liu & Hu, 1962 should be placed into the synonymy of Theloderma asperum (Boulenger, 1886), and that Philautus rhododiscus Liu & Hu, 1962 should be assigned to Theloderma , so as to correct the paraphyly. Additionally, the monophyly of ' Aquixalus ' is not supported, and this requires further examination. Results also indicate that the Rhacophorus leucomystax (Gravenhorst, 1829)/ Rhacophorus megacephalus (Hallowell, 1861) complex needs further revision. Studies employing broader sampling and more molecular markers will be needed to resolve the deep relationships within the subfamily Rhacophorinae.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 733–749.     

A new species of salamander of the genus Hynobius from Central Honshu, Japan (Amphibia, Urodela)

We describe a small salamander from south Central Honshu, Japan, as a new species, Hynobius katoi. The genetic distances between this species and several named species, including sympatric H. kimurae, derived from allozyme data from a starch gel electrophoresis, proved to be sufficiently large to differentiate it at a specific rank. Distribution of this species is confined to the montane regions of Shizuoka and Nagano Prefectures, on the Akaishi Mountains of the Chubu District, central Japan. It is regarded as a member of the naevius group of Hynobius, characterized by small number of large, pigmentless ova. The species differs from the other species of the naevius group by the combination of relatively small body size, nearly spotless body, relatively few vomerine teeth forming moderately shallow series, and unique electrophoretic pattern of isozymes.     

Desiccation resistance and water balance in southern African keratin beetles (Coleoptera, Trogidae): the influence of body size and habitat

Desiccation resistance and water balance were examined in the adults of seven trogid species, which differed both in body size and in the habitats from which they were collected. Body water contents (51–58% fresh mass) and desiccation rates at 27 °C (0.00026–0.00093 g h⁻¹) in these species were very similar to those of unrelated, similar-sized beetles from arid habitats. The keratin beetles differed markedly from many other adult Coleoptera by virtue of their very high haemolymph osmolality and inability to regulate haemolymph osmolality, and to catabolise lipids for water production, during desiccation. Like most other insects, the xeric trogid species had lower rates of water loss and longer survival times than trogids from mesic areas. This was due both to lower rates of water loss and to the larger body size of species from the more arid areas. Because absolute body water content was higher in large beetles than in small ones, larger body size conferred higher desiccation resistance on the very large Kalahari desert species. This suggests that there may be strong selection for large body size in such insects from arid areas. Most ecological and ecophysiological investigations of geographical variation in body size, and the species-body size distribution, have focused on temperature and metabolic rate as explanatory variables. This study suggests that attention should also be given to desiccation resistance. Accepted: 29 September 1997     

A new species of Platydracus Thomson, 1858 (Coleoptera,Staphylinidae, Staphylininae) from the upper Eocene Florissant beds,Colorado, USA

Platydracus breviantennatus n. sp., is described and figured based on an impression fossil from the upper Eocene Florissant beds of Colorado, the United States. Based on the large and densely setose body, relatively small eyes, and robust mandibles (right mandible seemingly with one simple preapical tooth), tibiae and antennae, the new species is placed in the modern genus Platydracus Thomson, 1858. It differs from other species of Platydracus by its large body, small head, and distinctly short antennae.     

Supplementary description and ontogenetic variability of Coryphaenoides affinis: rediscovery of the species after 145 years since its original description (Teleostei: Gadiformes: Macrouridae)

A large adult (138 mm head length) of the grenadier genus Coryphaenoides is described. It is identified as C. affinis hitherto known only from two juvenile type specimens collected by HMS Challenger off Uruguay in 1876. The diagnosis of the species is revised to include the discovery of considerable ontogenetic changes in squamation. Spinulation on the body scales are reduced or lost with size, while the armament of the head scales become amplified. The larger of the syntype specimens is designated as the lectotype. All three known specimens of C. affinis were collected in the southwestern Atlantic at depths between 3500 and 4000 m. A key for identification for the species of the subgenus Nematonurus is provided.     

Chromis okamurai,a new damselfish from the Okinawa Trough,Japan

A new pomacentrid fish,Chromis okamurai, is described from a single specimen taken from the Okinawa Trough (25°53.4′N, 123°59.4′E), in 135–175 m. This species is most closely related to the deep-dwellingC. mirationis, sharing with it the same meristic data (in particular XIV dorsal spines and 2 upper and lower caudal spinules), large eye, and relatively deep body. It differs in having a small round posterior nostril (large and slit like inmirationis), longer second anal spine, shorter pelvic fins, and in color: two dark brown stripes on body and a pale pectoral-fin axil (mirationis has a single, midlateral, yellow to yellowish brown stripe which extends broadly onto lower part of caudal fin, and a black pectoral axil).     

Allometries of the durations of torpid and euthermic intervals during mammalian hibernation: A test of the theory of metabolic control of the timing of changes in body temperature

Summary The durations of the intervals of torpor and euthermia during mammalian hibernation were found to be dependent on body mass. These relationships support the concept that the timing of body temperature changes is controlled by some metabolic process. Data were obtained from species spanning nearly three orders of magnitude in size, that were able to hibernate for over six months without food at 5°C. The timing of body temperature changes was determined from the records of copper-constantan thermocouples placed directly underneath each animal. Because all species underwent seasonal changes in their patterns of hibernation, animals were compared in midwinter when the duration of euthermic intervals was short and relatively constant and when the duration of torpid intervals was at its longest. Large hibernators remained euthermic longer than small hibernators (Fig. 2). This was true among and within species. The duration of euthermic intervals increased with mass at the same rate (mass^0.38) that mass-specific rates of euthermic metabolism decrease, suggesting that hibernators remain at high body temperatures until a fixed amount of metabolism has been completed. These data are consistent with the theory that each interval of euthermia is necessary to restore some metabolic imbalance that developed during the previous bout of torpor. In addition, small species remained torpid for longer intervals, than large species (Fig. 3). The absolute differences between different-sized species were large, but, on a proportional basis, they were comparatively slight. Mass-specific rates of metabolism during torpor also appear to be much less dependent on body mass than those during euthermia, but the precision of these metabolic measurements is insufficient for them to provide a conclusive test of the metabolic theory. Finally, small species with high mass-specific rates of euthermic metabolism are under tighter energetic constraints during dormancy than large species. The data presented here show that, in midwinter, small species compensate both by spending less time at high body temperatures following each arousal episode and by arousing less frequently, although the former is far more important energetically than the latter.     

Body size, biomic specialization and range size of African large mammals

Aim The goal of this paper is to examine the relationships between body size, biomic specialization and range size in the African large mammals, which are defined as all the African species corresponding to the orders Primates, Carnivora, Proboscidea, Perissodactyla, Hyracoidea, Tubulidentata, Artiodactyla and Pholidota. Location The study used the large mammal assemblage from Africa. Methods The degree of biomic specialization of African large mammals is investigated using the biomic specialization index (BSI) for each mammal species, based on the number of biomes it inhabits. Range size for each species is measured as the latitudinal extent of the geographical distribution of the species. We have analysed our data using both conventional cross‐species analyses and phylogenetically independent contrasts. Results There is a polygonal relationship between species biomic specialization and body size. While small and large species are biomic specialists, medium‐sized species are distributed along the whole range of biomic specialization. The latitudinal extent–body size relationship is approximately triangular. Small‐bodied species may have either large or small ranges, whereas large‐bodied ones have only large ranges. A positive correlation between latitudinal extent and biomic specialization is evident, although their relationship is better described as triangular. Main conclusions We found a polygonal relationship between species biomic specialization and body size, which agrees with previous arguments that small‐bodied species have more limited dispersal and, therefore, they may come to occupy a lesser proportion of their potential inhabitable biomes. On the other hand, large‐bodied species are constrained to inhabit biomes with a high productivity. A polygonal relationship between species latitudinal extent and body size in African large mammals agrees with previous studies of the relationship between range size and body size in other continents. The independent study of the macroecological pattern in biomic specialization highlights different factors that influence the body size–range size relationship. Although body size is usually implicated as a correlate of both specialization and geographical range size in large mammals, much of the variation in these variables cannot be attributed to size differences but to biome specific factors such as productivity, area, history, etc.     

Digenetic trematodes of the genus Clinostomum Leidy, 1856 (Digenea: Clinostomidae) from birds of Queensland, Australia, including C. wilsoni n. sp. from Egretta intermedia

Two species of Clinostomum previously described from Australia, C. hornum from Botaurus poiciloptilus (Australian bittern) and Nycticorax caledonicus (Nankeen night heron) and C. australiense from Pelecanus conspicillatus (Australian pelican), which have previously been synonymised with C. complanatum, are redescribed and recognised as valid species. In addition, C. complanatum is recorded from Egretta alba (large egret), E. garzetta (little egret), E. intermedia (plumed egret), N. caledonicus and Ardea novaehollandiae (white-faced heron). C. wilsoni n. sp. is described from E. intermedia from Queensland. C. wilsoni differs from the other three species in size and shape of the body and in the oral collar, oral sucker, intestinal caeca, caecal diverticula and position of testes. Taxonomic problems within the genus Clinostomum are discussed.