Molecular phylogeny,biogeography and character evolution of the montane genus Incarvillea Juss.(Bignoniaceae)

The complex orogeny of the Himalaya and the Qinghai-Tibet Plateau(QTP)fosters habitat fragmentation that drives morphological differentiation of mountain plant species.Consequently,determining phylogenetic relationships between plant subgenera using morphological characters is unreliable.Therefore,we used both molecular phylogeny and historical biogeographic analysis to infer the ancestral states of several vegetative and reproductive characters of the montane genus Incarvillea.We determined the taxonomic position of the genus Incarvillea within its family and inferred the biogeographical origin of taxa through Bayesian inference(BI),maximum likelihood(ML)and maximum parsimony(MP)analyses using three molecular data sets(trnL-trnF sequences,nr ITS sequences,and a data set of combined sequences)derived from 81%of the total species of the genus Incarvillea.Within the genus-level phylogenetic framework,we examined the character evolution of 10 key morphological characters,and inferred the ancestral area and biogeographical history of the genus.Our analyses revealed that the genus Incarvillea is monophyletic and originated in Central Asia during mid-Oligocene ca.29.42 Ma.The earliest diverging lineages were subsequently split into theWestern Himalaya and Sino-Himalaya during the early Miocene ca.21.12 Ma.These lineages resulted in five re-circumscribed subgenera(Amphicome,Olgaea,Niedzwedzkia,Incarvillea,and Pteroscleris).Moreover,character mapping revealed the ancestral character states of the genus Incarvillea(e.g.,suffruticose habit,cylindrical capsule shape,subligneous capsule texture,absence of capsule wing,and loculicidal capsule dehiscence)that are retained at the earliest diverging ancestral nodes across the genus.Our phylogenetic tree of the genus Incarvillea differs from previously proposed phylogenies,thereby recommending the placement of the subgenus Niedzwedzkia close to the subgenus Incarvillea and maintaining two main divergent lineages.

Genetic Diversity of MHC Gene and Molecular Phylogeny in Wanbei Cattle

[Objective] To investigate the genetic diversity, molecular phylogeny and origination in Wanbei cattle. [Methedl Blood samples were collected from 50 Wanbei cattle and 14 local cattle breeds. According to the bovine MHC gene sequence, one pair of primers was designed and used for PCR amplification. The sequences of amplified MHC genes were aligned to detect MHC haplotypes and nucleotide polyrnorphisms. A phy- Iogenetic tree based on the MHC gene was also constructed. [Result] A total of 24 polymorphic loci covering 7.86% of the entire length of the se- quences and seven MHC haplotypes were detected. Among these polymorphic loci, there were 18 transitions, 1 tranversions and 5 transition / tran- version. The difference in nucleotide sequence was 0.26% -4.53% between the haplotypes. The nucleotide diversity （π value） and haplotype di- versity （H） were 0.012 - 0.019 and 0.602 - 0.617, respectively. [ Conclusion ] Wanbei cattle do not have rich MHC genetic diversity. They have closer aenetic relationship with Luxi cattle, and both are within the same sister group.

Phylogeny and Geographical Derivation of Genus Hildenbrandia Based on Morphological and Molecular Evidence

The marine and freshwater Hildenbrandia samples were collected from diverse locations of China and Japan,and mor-phological characterization and molecular phylogenetic analysis were conducted on these samples.Morphological measurements of freshwater specimens were consistent with the results of H.jigongshanensis,whereas cell dimensions of marine specimens were slightly larger than those of widely distributed H.rubra and H.crouanii.Phylogenetic trees based on rbcL,psbA and UPA sequences were consistent.Freshwater specimens collected in this study formed an independent clade with H.jigongshanensis supported by high values.Species attribution was confirmed further by the similar ITS2 secondary structures among the samples of H.jigongshan-ensis.It was found that the intraspecific divergence of rbcL gene in H.jigongshanensis was smaller than other two freshwater species H.rivularis and H.angolensis.Phylogenetic trees showed that marine specimens in this study grouped together with H.rubra samples of North America,and were different with the H.rubra samples of Europe.Combining the comparison results of CBC numbers in ITS2 secondary structure,we propose the marine samples collected in this study to be a new species Hildenbrandia qingdaoensis sp.nov.The separation of the marine and freshwater specimens was supported by the rbcL,UPA and psbA phylogeny as well as ITS2 se-condary structure inference.Biogeographical reconstruction showed the ancestor of freshwater Hildenbrandia might derive from South America and dispersed from North America to Europe and then to Asian countries,which needs to be verified further with more sam-ples and molecular evidences from South America.

Morphological and Molecular Data Revealed One New Species of the Short-legged Toads Brachytarsophrys Tian and Hu,1983(Anura,Megophryidae)from Yunnan,China

A new species of the genus Brachytarsophrys,named Brachytarsophrys wenshanensis sp.nov.,has been identified in southeastern Yunnan,China.This new species can be readily distinguished from other known congeners by both morphological criterion and molecular analysis of three mitochondrial gene segments:16S,COI,and Cytb.This classification is based on the following morphological characters:(1)medium body size(SVL 83.8–85.1 mm in two adult males);(2)enormous head,with head width nearly 1.2 times the length;(3)tongue pyriform,feebly notched;(4)non-meeting heels;(5)male lacking nuptial pad;(6)tibiotarsal articulation reaching angle of mouth when hindlimbs are extended forward alongside the body;(7)absence of outer metatarsal tubercle,inner metatarsal tubercle elliptic and approximately equal to first toe;(8)rudimentary toe webbing,webbing formula:Ⅰ(2–)–(2^(++))Ⅱ(2^(–))–(3^(++))Ⅲ(2^(½))–(4)Ⅳ(4^(+))–(2^(⅔))V;(9)lateral fringes narrow;(10)dermal ridge or glandular fold on dorsum absent;(11)pectoral glands distinct and irregular,femoral gland small.Our work increases the number of species within the genus Brachytarsophrys to 9.

DNA barcoding of genus Toxoptera Koch （Hemiptera： Aphididae） Identification and molecular phylogeny inferred from mitochondrial COl sequences

Identification of aphid species is always difficult due to the shortage of easily distinguishable morphological characters. Aphid genus Toxoptera consists of species with similar morphology and similar to Aphis in most morphological characters except the stridulatory apparatus. DNA barcodes with 1 145 bp sequences of partial mitoehondrial cytochrome-coxidase I （COI） genes were used for accurate identification of Toxoptera. Results indicated mean intraspecific sequence divergences were 1.33%, whereas mean interspecific divergences were greater at 8.29% （0.13% and 7.79% if T. aurantii 3 and T. aurantii 4 are cryptic species）. Sixteen samples were distinguished to four species correctly by COI barcodes, which implied that DNA barcoding was successful in discrimination of aphid species with similar morphology. Phylogenetic relationships among species of this genus were tested based on this portion of COI sequences. Four species of Toxoptera assembled a clade with low support in maximum-parsimony （MP） analysis, maximumlikelihood （ML） analysis and Bayesian phylogenetic trees, the genus Toxoptera was not monophyletic, and there were two sister groups, such as T. citricidus and T. victoriae, and two clades of T. aurantii which probably presented cryptic species in the genus.

Molecular phylogeny of Pamphagidae （Acridoidea, Orthoptera） from China based on mitochondrial cytochrome oxidase II sequences

Phylogenetic relationships of Pamphagidae were examined using cytochrome oxidase subunit II （COIl） mtDNA sequences （684 bp）. Twenty-seven species of Acridoidea from 20 genera were sequenced to obtain mtDNA data, along with four species from the GenBank nucleotide database. The purpose of this study was analyzing the phylogenetic relationships among subfamilies within Pamphagidae and interpreting the phylogenetic position of this family within the Acridoidea superfamily. Phylogenetic trees were reconstructed using neighbor-joining （NJ）, maximurn parsimony （MP） and Bayesian inference （BI） methods. The 684 bp analyzed fragment included 126 parsimony informative sites. Sequences diverged 1.0%-11.1% between genera within subfamilies, and 8.8%-12.3% between subfamilies. Amino acid sequence diverged 0-6.1% between genera within subfamilies, and 0.4%-7.5% between subfamilies. Our phylogenetic trees revealed the monophyly of Pamphagidae and three distinct major groups within this family. Moreover, several well supported and stable clades were found in Pamphagidae. The global clustering results were similar to that obtained through classical morphological classification： Prionotropisinae, Thrinchinae and Pamphaginae were monophyletic groups. However, the current genus Filchnerella （Prionotropisinae） was not a monophyletic group and the genus Asiotmethis （Prionotropisinae） was a sister group of the genus Thrinchus （Thrinchinae）. Further molec- ular and morphological studies are required to clarify the phylogenetic relationships of the genera Filchnerella and Asiotmethis.

Identification, pathogenicity, and fungicide sensitivity of Eutiarosporella dactylidis associated with leaf blight on maize in China

Maize(Zea mays L.) is an economically vital grain crop that is cultivated worldwide. In 2011, a maize foliar disease was detected in Lingtai and Lintao counties in Gansu Province, China. The characteristic signs and symptoms of this disease include irregular chlorotic lesions on the tips and edges of infected leaves and black punctate fruiting bodies in dead leaf tissues. Given favourable environmental conditions, this disease spread to areas surrounding Gansu. In this study, infected leaves were collected from Gansu and Ningxia Hui Autonomous Region between 2018and 2020 to identify the disease-causing pathogen. Based on morphological features, pathogenicity tests, and multilocus phylogenetic analysis involving internal transcribed spacer(ITS), 18S small subunit rDNA(SSU), 28S large subunit rDNA(LSU), translation elongation factor 1-alpha(TEF), and β-tubulin(TUB) sequences, Eutiarosporella dactylidis was identified as the causative pathogen of this newly discovered leaf blight. Furthermore, an in vitro bioassay was conducted on representative strains using six fungicides, and both fludioxonil and carbendazim were found to significantly inhibit the mycelial growth of E. dactylidis. The results of this study provide a reference for the detection and management of Eutiarosporella leaf blight.

Taxonomy of Two New Peritrich Ciliates(Protozoa:Ciliophora)from Inland Saltwaters of China

The inland saltwater lakes harbor exceptional biodiversity.Here,two new species of solitary sessile peritrich ciliates were isolated from Qinghai Lake,the largest inland saltwater lake in China.Their morphology,ciliature,silverline system,and molecular phylogeny were investigated based on live observation,silver staining,and analysis of the small subunit ribosomal DNA(SSU rDNA).Vorticella paraglobosa sp.n.is characterized mainly by its obconical or elongate bell-shaped zooid,C-shaped macronucleus,single ventrally located contractile vacuole,two-rowed infundibular polykinety 3,and 28-38 silverlines between peristome and aboral tro-chal band and 10-15 between aboral trochal band and scopula.Vorticella cotyliformis sp.n.differs from its congeners mainly by its double-layered peristomial lip,cup-shaped zooid,J-shaped macronucleus,single ventrally positioned contractile vacuole,three-rowed infundibular polykinety 3,and 70-85 silverlines between peristome and aboral trochal band and 21-25 between aboral trochal band and scopula.The SSU rDNA sequences of the two new species were obtained,and the subsequent molecular phylogenetic analysis supported their taxonomic classification.

A New Species of the Genus Boulenophrys(Anura,Megophryidae)from Eastern Guangdong,China

A new horned toad species,Boulenophrys elongata sp.nov.,is described from Mt.Lianhua,Huidong County,eastern Guangdong,China.The new species is phylogenetically closest to B.brachykolos,which is restricted to the east of the Pearl River Estuary(i.e.,Shenzhen City and Hong Kong SAR).The new species can be distinguished from all recognized congeners by a combination of morphological characters and genetic divergences in the combined mitochondrial 16S,12S and CO1 genes.We recommend the new species be listed as Data Deficient(DD)in the IUCN categorization,pending further investigation.

A new genus of Asiatic short-tailed shrew (Soricidae, Eulipotyphla) based on molecular and morphological comparisons

Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. Blarinelline shrews were once widely distributed throughout Eurasia and North America, though only members of the Asiatic short-tailed shrew genus Blarinella currently persist （mostly in southwestern China and adjacent areas）. Only three forms of Blarinella have been recognized as either species or subspecies However, recent molecular studies indicated a strikingly deep divergence within the genus, implying the existence of a distinct genus-level lineage. We sequenced the complete mitochondrial genomes and one nuclear gene of three Asiatic short-tailed and two North American shrews and analyzed them morphometrically and morphologically. Our molecular analyses revealed that specimens ascribed to B. griselda formed two deeply diverged lineages, one a close relative to B. quadraticauda, whereas the other--comprised of topotype specimens from southern Gansu-iverged from other Blarinella in the middle Miocene （ca. 18.2 million years ago （Ma）, 95% confidence interval=13.4-23.6 Ma）. Although the skulls were similarly shaped in both lineages, we observed several diagnostic characteristics, including the shape of the upper p4. In consideration of the molecular and morphological evidence, we recognize B. griselda as the sole species of a new genus, namely, Pantherina gen. nov. Interestingly, some characteristics of Pantherina griselda are more similar to fossil genera, suggesting it represents an evolutionarily more primitive form than Blarinella.Recognition of this new genus sheds light on the systematics and evolutionary history of the tribe Blarinellini throughout Eurasia and North America.

Phylogeny of Ptychostomum (Bryaceae,Musci) inferred from sequences of nuclear ribosomal DNA internal transcribed spacer (ITS) and chloroplast rps4

The phylogeny of Ptychostomum was first spacer （ITS） region of the nuclear ribosomal （nr） DNA DNA rps4 sequences. Maximum parsimony, maximum undertaken based on analysis of the internal transcribed and by combining data from nrDNA ITS and chloroplast likelihood, and Bayesian analyses all support the conclusion that the reinstated genus Ptychostomum is not monophyletic. Ptychostomum funkii （Schwagr.） J. R. Spence （≡ Bryum funkii Schwaigr.） is placed within a clade containing the type species of Bryum, B. argenteum Hedw. The remaining members of Ptychostomum investigated in the present study constitute another well-supported clade. The results are congruent with previous molecular analyses. On the basis of phylogenetic evidence, we agree with transferring B. amblyodon Mull. Hal. （≡ B. inclinatum （Brid.） Turton≡ Bryum archangelicum Bruch ＆ Schimp.）, Bryum lonchocaulon Mull. Hal., Bryum pallescens Schleich. ex Schwaigr., and Bryum pallens Sw. to Ptychostomum.

Phylogeny of Limenitidinae Butterflies (Lepidoptera: Nymphalidae) Inferred from Mitochondrial Cytochrome Oxidase I Gene Sequences

The phylogenetic analyses of the subfamily Limenitidinae are performed based on 1 471 bp of mtDNA cytochrome oxidase subunit I （COI） gene sequence data which were obtained from 21 individuals spanning 9 genera, along with those of 17 species obtained from GenBank, using Apatura iris, Aglais urticae, and Polyura dolon as outgroup species. Although the transitions at the third codon positions of the COI data set were highly saturated, they were still retained for analysis as they contain the majority of the phylogenetic information, and thus, the maximum pasimony （MP） under different weighting schemes and maximum likelihood （ML） trees were reconstructed in this study. The results showed that within this subfamily, the results based on the COI gene sequences are approximately identical to the traditional classification results. However, the clustering of Lexias pardalis and Tanaecia julii within the genus Euthalia as well as the clustering of Phaedyma aspasia within the genus Neptis with weak support are different from that of the current classification scheme made by Chinese scholars. The genus Limenitis is splited into two subclusters in the trees constructed by using MP and ML methods. These results support one of the strongest hypotheses for the tribe relationships within Limenitidinae.

A revision of Dryopteris sect.Diclisodon(Dryopteridaceae)based on morphological and molecular evidence with description of a new species

Dryopteris sect.Diclisodon is a small section of ferns with about 12 species mainly distributed in East Asia.Here,we carried out morphological and phylogenetic analyses of this section.A new species from southwest China,D.gaoligongensis,is described and illustrated.Dryopteris gaoligongensis resembles D.indonesiana and D.sparsa,but differs by having a creeping rhizome and large 4-pinnate fronds.We also show that D.glabrior Ching&Z.Y.Liu is a distinct species;however,because it is a later homonym of D.glabrior Copel.,it should be renamed D.renchangiana.We conclude that a species previously known as D.nitidula,also an illegitimate homonym,should be recognized with a new name,D.sinonepalensis.We resolve the phylogenetic position of D.yoroii as sister to other sampled species of D.sect Diclisodon.Our phylogenetic analyses confirm the distinctiveness of D.gaoligongensis,D.renchangiana,and D.sinonepalensis.A key to species of D.sect Diclisodon in China is provided.

Detection and Molecular Characterization of Enteroviruses in Korean Surface Water by Using Integrated Cell Culture Multiplex RT-PCR

Objective To identify waterborne enteric viruses in Korean surface water. Methods Integrated cell culture（ICC）multiplex reverse transcription-polymerase chain reaction （RT-PCR） was simultaneously designed to detect coxsackieviruses （CV）, polioviruses （PV）, and reoviruses （RV）. ICC-multiplex RT-PCR and phylogenetic analysis were conducted using 21 total culturable virus assay （TCVA）-positive sample-inoculated cell cultures. Results CV and RV were detected in 9 samples each, and 3 samples were positive for both CV and RV. PV was not detected in any sample. Molecular phylogenetic analysis of the VP1 gene sequences revealed that CV types B2 and B4 predominated in Korean surface water, and the nucleotide sequences of CV type B2 were clustered with those of CVs isolated from China and Japan. The results suggested that the evolution of these viruses occurred in a region-specific manner. Conclusion CV and RV are detectable in Korean surface water, with a predominance of CV type B2, and the evolution of CV type B2 occur in a region-specific manner.

Molecular Phylogenetic Analysis of the Main Lineages of Nymphalinae(Nymphalidae: Lepidoptera) Based on the Partial Mitochondrial COI Gene

The phylogenetic relationships of the subfamily Nymphalinae （sensu Chou 1994） were analyzed based on 1 488 bp of mtDNA cytochrome oxidase subunit I （COI） gene sequence data obtained from 24 individuals, along with those of eight species obtained from GenBank. The base compositions of this COI fragment varied among the individuals as follows： T 39.9%, C 14.6%, A 32.2%, and G 13.4%, with a strong AT bias （72.1%）, as usually found in insect mitochondrial genomes. The A ＋T contents of the third, second, and first codon positions of the COI fragments in this study was 92.4, 62.2, and 61.4%, respectively. The phylogenetic trees were reconstructed by neighbor-joining （NJ）, maximum likelihood （ML）, and Bayesian methods by using Byblia anvatara as outgroup. Phylogenetic analyses based on the COI gene sequence data created very similar topologies, which were producing trees with two main clades A and B, and five subclades. The data indicated that the tribes Nymphalini and Hypolimni （sensu Chou 1994） are not monophyletic groups, and the genus Junonia should be removed from Nymphalini to Hypolimni （=Junoniini）. On the basis of the data, the Symbrenthia and Araschnia had a relative distant relationship with the rest of Nymphalini. The relationships of species in the Nymphalini were confirmed via the NJ, ML, and Bayesian methods, namely （（（（Nymphalis ＋ Kaniska） ＋ Polygonia） ＋Aglais） ＋ Vanessa） ＋ （Symbrenthia ＋Araschnia）. This investigation provides a little novel information for Chinese researches of butterflies.

A New Species of the Genus Hebius Thompson,1913(Serpentes:Natricidae)from Yunnan,China

A new keelback snake of the genus Hebius,H.jingdongensis sp.nov.,is reported from Jingdong in Yunnan,China based on a combination of molecular and morphological evidence.Although closest to H.khasiensis and H.boulengeri phylogenetically,this new species is genetically distinct amongst congeners with the mitochondrial Cyt b uncorrected p-distance ranging from 9.5%(vs.H.octolineatus)to 14.8%(vs.H.andreae).In addition,this new species can be distinguished from its congeners by a combination of the following morphological characters:(1)body cylindrical,tail relatively short,TaL/TL 0.235–0.243;(2)dorsal scale rows 19-19-17;(3)ventrals 163–166,cloacal plate divided,subcaudals 71–74 pairs;(4)supralabials 8,the first two contact with the nasals;(5)infralabials 9 or 10,preocular 1,postoculars 3;(6)body dorsum sandy beige with the sides yellowish-brown decorated with two brick-red stripes on the dorsolateral scale rows in life;(7)sutures between the labials brick-red in life and taupe in preservative;(8)ventral light yellow with the outer part of each scale brick-red in life;(9)ventrolateral stripes present.

Plastid phylogenomics and species discrimination in the“Chinese”clade of Roscoea(Zingiberaceae)

Roscoea is an alpine or subalpine genus from the pan-tropical family Zingiberaceae,which consists of two disjunct groups in geography,namely the"Chinese"clade and the"Himalayan"clade.Despite extensive research on the genus,Roscoea species remain poorly defined and relationships between these species are not well resolved.In this study,we used plastid genomes of nine species and one variety to resolve phylogenetic relationships within the"Chinese"clade of Roscoea and as DNA super barcodes for species discrimination.We found that Roscoea plastid genomes ranged in length from 163,063 to 163,796 bp,and encoded 113 genes,including 79 protein-coding genes,30 tRNA genes,four rRNA genes.In addition,expansion and contraction of the IR regions showed obvious infraspecifc conservatism and interspecific differentiation.Plastid phylogenomics revealed that species belonging to the"Chinese"clade of Roscoea can be divided into four distinct subclades.Furthermore,our analysis supported the independence of R.cautleoides var.pubescens,the recovery of Roscoea pubescens Z.Y.Zhu,and a close relationship between R.humeana and R.cautloides.When we used the plastid genome as a super barcode,we found that it possessed strong discriminatory power(90%)with high support values.Intergenic regions provided similar resolution,which was much better than that of protein-coding regions,hypervariable regions,and DNA universal barcodes.However,plastid genomes could not completely resolve Roscoea phylogeny or definitively discriminate species.These limitations are likely related to the complex history of Roscoea speciation,poorly defined species within the genus,and the maternal inheritance of plastid genomes.

A New Species of Theloderma Tschudi,1838(Amphibia:Rhacoporidae)from Central Java Allied to T.horridum(Boulenger,1903)

Species diversity of the genus Theloderma has been extensively studied in recent decades,with the majority of newly described species hailing from the Indochina region.We describe a new species of mediumsized Theloderma based on molecular and morphological evidence from lowland forest in the southern part of Central Java Province,Indonesia,that is highly similar to and considered cryptic within the T.horridum group.A phylogenetic approach shows that this population is a distinct genetic lineage and can be distinguished from its congeners by the absence of vomerine teeth,moderate irregular blunt warts and studded with calcified asperities on the dorsal skin,a distinct color pattern,and a distinct denticle formula in the tadpole.Although the range of the new species is known only from Central Java,it cannot be ruled out that it may extends into adjacent areas,such as other parts of Java and Sumatra.

Tibet,the Himalaya,Asian monsoons and biodiversity-In what ways are they related？

Prevailing dogma asserts that the uplift of Tibet, the onset of the Asian monsoon system and high biodiversity in southern Asia are linked, and that all occurred after 23 million years ago in the Neogene.Here, spanning the last 60 million years of Earth history, the geological, climatological and palaeontological evidence for this linkage is reviewed. The principal conclusions are that: 1) A proto-Tibetan highland existed well before the Neogene and that an Andean type topography with surface elevations of at least 4.5 km existed at the start of the Eocene, before final closure of the Tethys Ocean that separated India from Eurasia. 2) The Himalaya were formed not at the start of the India-Eurasia collision, but after much of Tibet had achieved its present elevation. The Himalaya built against a pre-existing proto-Tibetan highland and only projected above the average height of the plateau after approximately 15 Ma. 3)Monsoon climates have existed across southern Asia for the whole of the Cenozoic, and probably for a lot longer, but that they were of the kind generated by seasonal migrations of the Inter-tropical Convergence Zone. 4) The projection of the High Himalaya above the Tibetan Plateau at about 15 Ma coincides with the development of the modern South Asia Monsoon. 5) The East Asia monsoon became established in its present form about the same time as a consequence of topographic changes in northern Tibet and elsewhere in Asia, the loss of moisture sources in the Asian interior and the development of a strong winter Siberian high as global temperatures declined. 6) New radiometric dates of palaeontological finds point to southern Asia's high biodiversity originating in the Paleogene, not the Neogene.

A New Species of the Genus Tylototriton(Amphibia,Salamandridae) from Hunan,China

A new species of the genus Tylototriton is described in this article based on morphological and genetic evidence. The new species is identified belonging to the T. asperrimus group and is similar to T. wenxianensis. The diagnostic characters of the new species are as follows: dorsal ridge broad and thick,its width approximately equal to eye diameter(P < 0.001); tail height greater than width at base of tail(P < 0.001); no villous genital papilla found inside the male anal fissure; nodule-like warts,along lateral margin of the trunk,bulge and forming tubercles,and thin and transverse striae present between the tubercles.