A new megophryid species is described from southwestern Sarawak, Malaysian Borneo. In appearance, Leptolalax marmoratus sp. nov. is most similar to L. hamidi also from southwestern Sarawak, but differs from it by mtDNA sequence, larger body size, and higher dominant frequency of advertisement call. The assumption that more than one species of Leptolalax coexist at one locality in Borneo is supported. The finding of the new species raises the species number of Leptolalax known from Borneo to nine, and the island is thought to be one of the diversification centers of the genus.
We record a tree frog of the genus Chiromantis for the first time from outside the Southeast Asian continent and describe it as a new species, Chiromantis inexpectatus. The new species from the Malaysian state of Sabah, Borneo, is a small-sized Chiromantis (male snout-vent length ca. 22 mm), and is distinguished from all other members of the genus by the combination of the following morphological characteristics: dark stripes absent, but dark spots present on dorsum; a dark-brown lateral band present from snout tip to half of body, bordered ventrally by white stripe; third and fourth fingers less than half webbed; third finger disk wider than tympanum diameter; and inner metatarsal tubercle present. Significance of findings of this species from Borneo Island, as well as phylogeny and breeding habit of the genus Chiromantis, are briefly discussed.
In general, African catfish shows higher survival rates in the dark conditions than in the light conditions. In this study, larval behavior of African catfish was observed under 0, 0.01, 0.1, 1, 10, and 100 lx using a CCD camera to investigate the reason why African catfish larvae show higher survival rates in dark conditions. The larvae showed significantly higher swimming activity under 0, 0.01, and 0.1 lx than that under 10 and 100 lx. The larvae also showed significantly increased aggressive behavior under 10 and 100 lx; the swimming larvae attacked resting individuals more frequently under 10 and 100 lx than under 0, 0.01, and 0.1 lx. The aggressive behavior and sharp teeth of the attacking larvae appeared to induce skin surface lesions on injured larvae. Chemical substances were then generated from the injured skin surface, and these chemical stimuli triggered cannibalistic behavior in other fish near the injured fish. The results of this study demonstrate that the higher survival rates of African catfish larvae under dark conditions are a result of inactivity and subsequent increase in chemical releasing stimuli concentrations around inactive individuals that triggers feeding behavior in nearby active catfish. Therefore, we recommend larval rearing of African catfish in dark or dim conditions, as it improves catfish survival rates.
Although the crab-eating frog Fejervarya cancrivora is one of the most widely distributed species in Asian region, taxonomic relationships among different populations remain unclarified. In this study, we attempted to elucidate the taxonomic status of F. cancrivora from Indonesian and other Asian populations. Five populations of F. cancrivora from Selangor (Malaysia), Cianjur (Java, Indonesia), Trat (Thailand), Khulna (Bangladesh), and Makassar (Sulawesi, Indonesia) were morphologically observed and subjected to crossing experiments. Principal component and clustering analyses revealed that these five populations could be organized into three groups corresponding to three observed morphological types: a Selangor and Cianjur group (large-type), a Trat and Khulna group (mangrove-type), and a Makassar group (Sulawesi-type). The limited crossing experiments revealed that hybrids between Selangor females and Cianjur and Trat males developed normally, whereas hybrids between Selangor females and Khulna males showed incomplete gametic isolation. Histological observations of the testes of mature males revealed the presence of pycnotic nuclei in the hybrids between Selangor females and Khulna males in addition to normal bundles of spermatozoa. In contrast, no pycnotic nuclei were observed in the Selangor controls. Although meiotic metaphases in the controls were normal, those in hybrids showed several abnormalities, such as the appearance of univalents and an increase in rod-shaped bivalents. Based on our findings from the morphological observations and crossing experiments, we conclude that each of three identified types represents a distinct species. We propose that the large-type is F. cancrivora, the mangrove-type is F. moodiei, and the Sulawesi-type represents an undescribed species.
A new microhylid, Kalophrynus yongi, Is described from the Cameron Highlands of Peninsular Malaysia. Morphologically, the new species differs from all known congeners by having a very stout forelimb with a humeral spine in males. Acoustically, it resembles K. baluensis and K. heterochirus and sharply differs from K. interlineatus, K. pleurostigma, K. palmatissimus, and K. nubicola.
A new species in the rhacophorid genus Philautus is described on the basis of five male specimens collected from the Matang Range, a herpetologically well-surveyed area of the lowland of Sarawak, western Borneo. The species possesses a cutaneous pectoris muscle but lacks vomerine teeth, and is assigned to the aurifasciatus group of Philautus. It is superficially similar to some other species of the same group, but can be differentiated from them by a combination of several morphological characters, including lack of a nuptial pad and lingual papilla, and more significantly, by a distinct advertisement call. This discovery underscores the immediate necessity for detailed surveys in the already well-explored lowlands of Borneo.
A new megophryid species, Leptolalax kecil, is described from the Cameron Highlands of Peninsular Malaysia on the basis of acoustic and morphological characteristics. It has an advertisement call dissimilar to that of other, congeneric species, and is the smallest of the known Leptolalax. It is similar to L. pluvialis from Vietnam in small body size, but differs from it in ventral color, and in the size and color of the pectoral gland. The distributional pattern of the new species Is discussed.
To elucidate genetic divergence and evolutionary relationship in Fejervarya cancrivora from Indonesia and other Asian countries, allozyme and molecular analyses were carried out using 131 frogs collected from 24 populations in Indonesia, Thailand, Bangladesh, Malaysia, and the Philippines. In the allozymic survey, seventeen enzymatic loci were examined for 92 frogs from eight representative localities. The results showed that F. cancrivora is subdivided into two main groups, the mangrove type and the large- plus Pelabuhan ratu types. The average Nel's genetic distance between the two groups was 0.535. Molecular phylogenetic trees based on nucleotide sequences of the 16S rRNA and Cyt b genes and constructed with the ML, MP, NJ, and BI methods also showed that the individuals of F. cancrivora analyzed comprised two clades, the mangrove type and the large plus Pelabuhan ratu / Sulawesi types, the latter further split into two subclades, the large type and the Pelabuhan ratu / Sulawesi type. The geographical distribution of individuals of the three F. cancrivora types was examined. Ten Individuals from Bangladesh, Thailand, and the Philippines represented the mangrove type; 34 Individuals from Malaysia and Indonesia represented the large type; and 11 individuals from Indonesia represented the Pelabuhan ratu / Sulawesi type. Average sequence divergences among the three types were 5.78-10.22% for the 16S and 12.88-16.38% for Cyt b. Our results suggest that each of the three types can be regarded as a distinct species.
We describe a new species of torrent-dwelling ranid frog of the genus Meristogenys from the Crocker Range, western Sabah, northern Borneo. The new species, Meristogenys maryatiae, differs from congeners by the combination of: small body, males 31-37 mm and females 65-66 mm in snout-vent length; head narrower than long; eyes moderate, diameter subequal to snout; iris unicolored; legs long; ventral surface of tibia without heavy pigmentation; rear of thigh blotched dark brown and cream; toes fully webbed; outer metatarsal tubercle present; larval dental formula 7(4-7)/6(1).
The present study was conducted to elucidate the genetic divergence and the phylogenetic relationships in the F. limnocharis complex from Bangladesh and other Asian countries such as Sri Lanka, Thailand, Malaysia, Taiwan and Japan by allozyme analyses. We used a total of 95 frogs of the F. limnocharis complex from these countries and F. cancrivora from the Philippines as an outgroup. Based on body size, the F. limnocharis complex from Bangladesh was divided into three distinct groups: large, medium and small types. Allozyme analyses were carried out with 28 loci encoding 20 enzymes and two blood proteins by horizontal starch-gel electrophoresis. When genetic distance was calculated, distinct divergence was found among the three types: mean genetic distance was 0.782 between the small and medium types, 1.458 between the large and medium types, and 1.520 between the large and small types. Phylogenetic trees based on genetic distance showed that all populations of Bangladesh small type strongly formed a cluster and were found to be most closely related to the Sri Lanka population; that all populations of Bangladesh large type formed a very strong cluster and were grouped with several populations from Thailand, Malaysia, Japan, and Taiwan; and that the medium type was segregated from all other groups. This may imply that each of the three types is a different species, and that the medium type is possibly an undescribed taxon.
Examination of types and recently collected specimens revealed that Ansonia anotis Inger, Tan, and Yambun, 2001 and Pedostibes maculatus (Mocquard, 1890), both described from Kinabalu, Sabah, Malaysia, are hardly differentiated morphologically. Analyses of a total of 2,427 bp of the 12S rRNA, tRNA(val), and 16S mitochondrial rRNA genes revealed that the two species are very close genetically. Thus A. anotis is regarded as conspecific and is synonymized with P. maculatus. Genetically, this species proved to form a lineage distinct from other bufonids from Southeast Asia, including species of Ansonia and Pedostibes. Because the species has also some unique morphological traits different from known bufonid genera, we propose to establish a new genus for Nectophryne maculata Mocquard, 1890.
In order to elucidate the taxonomic status of the Fejervarya limnocharis complex relative to Malaysia and Japan populations, morphological observations and molecular phylogenetic analysis were carried out using three populations from Indonesia (type locality), Malaysia, and Japan. In addition, we conducted histological and spermatogenic observations using hybrids among these populations. Principal component and cluster analyses demonstrated that these populations could be clearly separated from one another. Abnormal testes were found in the hybrids between the Japan and Indonesia populations and between the Japan and Malaysia populations, but testes of the controls and hybrids between the Malaysia and Indonesia populations were quite normal. The mean number of univalents per cell was 5.42, 4.58, and 0.20 in hybrids between the Indonesia and Japan populations, Malaysia and Japan populations, and Indonesia and Malaysia populations, respectively. Sequence divergences in 16S rRNA and Cyt b genes were 0-0.4% (xbar=0.2%) and 0.3-1.5% (xbar=1.0%), respectively, between the Malaysia and Indonesia populations, and 2.4-2.6% (xbar=2.5%) and 11.0-12.0% (xbar=11.5%) between the Japan population and F. limnocharis complex, including the Malaysia and Indonesia populations and F. multistriata from China. This study indicated that the Malaysia population and F. multistriata from China should be designated as a subspecies of topotypic F. limnocharis, and that the Japan population should be regarded as a distinct species.
This study was conducted to clarify the development of free neuromasts with growth of the barramundi, Lates calcarifer. A pair of free neuromasts was observed behind the unpigmented eyes in newly hatched eleutheroembryos with a mean total length of 1.93 mm, and two-hour-old eleuthero-embryos could respond to an approaching pipette. At 2 days after hatching, the egg yolk sac was mostly consumed, the eyes were pigmented, and the larvae commenced feeding on rotifers. Free neuromasts increased in number with growth and commenced developing into canal neuromasts in barramundi 15 days old with a mean total length of 8.07 mm. The average length of the major axis of the trunk free neuromasts attained approximately 12.9-15.5 microm, and the number of sensory cells was 15.4-17.5 at 15-20 days old. Developed cupulae of free neuromasts were observed in 1-day-old eleutheroembryos. The direction of maximum sensitivity of free neuromasts, determined from the polarity of the sensory cells, coincided with the minor axis of the lozenge-shaped outline of the apical surface of the free neuromasts. The polarity of trunk neuromasts was usually oriented along the antero-posterior axis of the fish body, but a few had a dorso-ventral direction. On the head, free neuromasts were oriented on lines tangential to concentric circles around the eye.
We examined allozyme variation in two camaenid tree snails, Amphidromus atricallosus and A. inversus, across two principal regions of Thailand and from Singapore, plus for A. inversus, one site in peninsular Malaysia. Using horizontal starch gel electrophoresis, 13 allozyme loci (11 polymorphic) were screened for A. atricallosus and 18 (5 polymorphic) for A. inversus. Heterozygosity was higher in A. atricallosus (Hexp=0.018-0.201, mean=0.085) than in A. inversus (Hexp=0-0.023, mean= 0.002). Genetic heterogeneity among samples was higher in A. inversus (Fst=0.965) than in A. atricallosus (Fst=0.781). Within A. atricallosus, populations were more differentiated in southern Thailand (Fst=0.551) than in eastern Thailand (Fst=0.144). The high Fst and low Hexp in populations of A. inversus suggest that this species is likely to have experienced a series of strong bottlenecks, perhaps occurring chiefly on offshore continental-shelf islands. The low Fst values of A. atricallosus in eastern Thailand suggest frequent gene flows among populations in this region. The southern and eastern samples of A. atricallosus exhibited fixed allele differences at four loci and great genetic distance (Nei's D=0.485-0.946), suggesting that these two samples may actually represent, or else be evolving into, separate species.
Two forms of Staurois that are differentiated by body size occur parapatrically in the Crocker Range, Sabah, Borneo. Analyses of a total of 1,499 bp of the mitochondrial cytochrome b, 12S rRNA, and 16S rRNA genes revealed that the two forms could be completely split genetically. The two forms could be also clearly differentiated morphologically, not only by snout-vent length but also by the relative sizes of snout, eye, and finger disk. Comparisons of the two forms with all known species of the genus revealed the large and small forms to be S. tuberilinguis and S. parvus, respectively. The latter species has long been synonymized with the former, but we here consider them to represent different species.
This study examined 396 base pairs of the mitochondrial cytochrome b gene from 110 individuals belonging to the genus Hampala, a group of freshwater cyprinids that inhabit Southeast Asia. The samples were taken from various locations throughout Sarawak, Sabah, and peninsular Malaysia. The nucleotide sequences were subjected to phylogenetic analyses by using the neighbor-joining, maximum parsimony, and maximum likelihood methods. All three methods revealed the reciprocally monophyletic relationship of Hampala macrolepidota to the other Hampala forms, thus strongly supporting its status as a distinct species. Phylogenetic analysis also discovered the existence of two H. bimaculata lineages endemic to Borneo: (1) a newly identified species from the southern and central part of Sarawak assigned as H. bimaculata Type A and (2) the previously described H. bimaculata from northern Sarawak and the west coast of Sabah assigned as H. bimaculata Type B. However, the status of H. sabana and an intermediate form were not elucidated. The results suggest that the intermediate form from the Tawau population is actually a subpopulation of H. sabana, while the highly divergent intermediate form from Kalabakan could represent a cryptic species. The sharing of H. macrolepidota haplotypes in the southern peninsular Malaysia and southern and central Sarawak samples (Hm1 and Hm2) reflected the recent disconnection of the two regions, during the late Pleistocene. Overall, the partial sequence of the mitochondrial cytochrome b gene was useful for resolving the phylogenetic relationships among Hampala fishes in Malaysia.
Three new megophryid species, Leptolalax melanoleucus, L. fuliginosus, and L. solus, are described from southwestern and southern Thailand on the bases of acoustic and morphological characteristics. Leptolalax melanoleucus and L. fuliginosus are similar to L. pelodytoides from northern Thailand, but differ from it completely in advertisement call characteristics and ventral color. Leptolalax solus is similar to L. heteropus from peninsular Malaysia, but differs from it by advertisement call, as well as by some body proportions. The distributional pattern of Leptolalax within Thailand is discussed.
We describe a new species of cascade frog of the genus Rana, from west Malaysia. Rana monjerai, new species is a medium-sized frog of the subgenus Odorrana (SVL of males, 38-43 mm; of one female, 75 mm), and is distinguished from all other members of this subgenus by the combination of: white lip stripe, dorsolateral fold, full web on the fourth toe, vomerine teeth, gular vocal pouch and relatively large tympanum in males, no dorsal marking, no clear light spots on rear of thigh, first finger subequal to second, finely tuberculated dorsum, and unpigmented ova. The significance of finding this species from peninsular Malaysia is discussed.
Mesohabitat selection in fluvial fishes was studied in a small tropical stream of the Malay Peninsula. A total of 681 individuals representing 24 species were sampled at 45 stations within heterogeneous stream (ca. 1 km in length), in which water depth, water velocity, substrate size, and riparian canopy cover were measured as environmental variables. A canonical correspondence analysis (CCA) yielded a diagram that shows a specific mesohabitat selection of the fish assemblage, in which the species were plotted widely on the CCA1-CCA2 biplot. Generalized linear model also revealed a significant pattern of the mesohabitat selection of several species. Water velocity and substrate size mainly separated on CCA1, indicating variation of pool (deep, slow-flow section) and riffle (shallow, fast-flow section) structures is a primary factor of mesohabitat selection in the fluvial fish assemblage. The mean body weight of species significantly correlated with CCA1; larger species tended to inhabit pools, while small ones occupied riffles. The riparian canopy cover separated on CCA2. The trophic level of species significantly correlated with CCA2; herbivorous species (low trophic level) selected open sites without riparian cover, whereas omnivorous/carnivorous (middle-high trophic level) species preferred highly covered sites. In conclusion, our results suggest that mesohabitat selection is closely related to the species feeding habit, which is consistent with the results of previous studies.
A cryptic Bornean torrent frog of the genus Meristogenys, which is divergent genetically and morphologically from all known congeners, is described from mountain streams of western Sarawak, East Malaysia (Borneo). The species occurs sympatrically with the type species of the genus, M. jerboa, but apparently differs from it in adult coloration and larval morphology, such as keratodont formulae and glands in tail fins. Females of the new species possess much larger and fewer eggs than in sympatric M. jerboa, suggesting significantly different reproductive traits between these species. A key to larvae of known species of the genus is provided.