Genomic clues to the parental origin of the wild flowering cherry <Emphasis Type="Italic">Prunus yedoensis</Emphasis> var. <Emphasis Type="Italic">nudiflora</Emphasis> (Rosaceae)

Prunus yedoensis Matsumura is one of the popular ornamental flowering cherry trees native to northeastern Asia, and its wild populations have only been found on Jeju Island, Korea. Previous studies suggested that wild P. yedoensis (P. yedoensis var. nudiflora) is a hybrid species; however, there is no solid evidence on its exact parental origin and genomic organization. In this study, we developed a total of 38 nuclear gene-based DNA markers that can be universally amplifiable in the Prunus species using 586 Prunus Conserved Orthologous Gene Set (Prunus COS). Using the Prunus COS markers, we investigated the genetic structure of wild P. yedoensis populations and evaluated the putative parental species of wild P. yedoensis. Population structure and phylogenetic analysis of 73 wild P. yedoensis accessions and 54 accessions of other Prunus species revealed that the wild P. yedoensis on Jeju Island is a natural homoploid hybrid. Sequence-level comparison of Prunus COS markers between species suggested that wild P. yedoensis might originate from a cross between maternal P. pendula f. ascendens and paternal P. jamasakura. Moreover, approximately 81% of the wild P. yedoensis accessions examined were likely F1 hybrids, whereas the remaining 19% were backcross hybrids resulting from additional asymmetric introgression of parental genotypes. These findings suggest that complex hybridization of the Prunus species on Jeju Island can produce a range of variable hybrid offspring. Overall, this study makes a significant contribution to address issues of the origin, nomenclature, and genetic relationship of ornamental P. yedoensis.     

Origins of Japanese flowering cherry (Prunus subgenus Cerasus) cultivars revealed using nuclear SSR markers

Japanese flowering cherry (Prunus subgenus Cerasus) cultivars, which are characterized by beautiful flowers, have been developed through hybridization among wild Prunus taxa. The long history of cultivation has caused significant confusion over the origins of these cultivars. We conducted molecular analysis using nuclear simple sequence repeat (SSR) polymorphisms to trace cultivar origins. Bayesian clustering based on the STRUCTURE analysis using SSR genotypes revealed that many cultivars originated from hybridization between two or more wild species. This suggests that morphological variations among flowering cherry cultivars probably arose through a complex sequence of hybridizations. Our findings generally supported estimates of the origins of cultivars based on morphological study, although there were some exceptions.     

Development of 13 EST-SSRs for Cerasus jamasakura and their transferability for Japanese flowering cherries

Simple sequence repeat (SSR) markers were developed for the flowering cherry Cerasus jamasakura (also known as Prunus jamasakura) using 31,995 expressed sequence tags (ESTs) from the NCBI database. Out of 96 of designed primer pairs, 63 showed clear PCR amplification and 13 of these revealed polymorphism in eight individuals sampled across the species’ range. The number of alleles detected and expected heterozygosity ranged from 1 to 8 and 0.000 to 0.833, respectively, when these 13 loci were examined in 23 individuals from a single population. For all except one of the lcoi, polymorphism was also detected in at least four of six other taxa of flowering cherries examined. The results show that the developed EST-SSRs are highly transferable, and that these markers are likely to be useful in studies of the population genetics of flowering cherries.     

Complete chloroplast genome of cultivated flowering cherry, <Emphasis Type="Italic">Prunus</Emphasis> ×<Emphasis Type="Italic">yedoensis</Emphasis> ‘Somei-yoshino’ in comparison with wild <Emphasis Type="Italic">Prunus yedoensis</Emphasis> Matsum. (Rosaceae)

Prunus ×yedoensis Matsum. ‘Somei-yoshino’ is the most common and widespread cultivar of the ornamental flowering cherries. We hereby report its complete chloroplast (cp) genome sequences generated by whole-genome next-generation sequencing approach. The cp genome size was 157,792 bp in length consisting of four regions; large single-copy region (85,914 bp), small single-copy region (19,120 bp), and a pair of inverted repeat regions (26,379 bp). The genome contained a total of 131 genes, including 86 coding genes, 8 rRNA genes, and 37 tRNA genes. A total of 92 simple sequence repeats (SSRs) were detected within the cp genome. Its molecular features were compared with the complete cp genome of wild P. yedoensis, which occurs rarely in natural habitats of Mt. Halla in Jeju Island, Korea, displaying nearly indistinguishable morphology as P. ×yedoensis ‘Somei-yoshino’. Although both cp genomes were structured highly alike, the sequence variations between them were revealed in several single-nucleotide polymorphisms (SNPs). Using additional individuals of wild and cultivated flowering cherries, PCR amplification confirmed that those SNPs were phylogenetically informative, providing distinction between wild and cultivated flowering cherries. In future study, the SNPs and SSRs reported in this study could be used to identify wild individuals from morphologically identical cultivars of flowering cherries and also to conserve the genetic diversity of wild flowering cherries in Jeju Island.     

THE HYBRID ORIGIN OF ASTER ASCENDENS (ASTERACEAE)

Aster ascendens Lindl. is a rhizomatous perennial of the Great Basin region of the western United States, with a chromosome base number of x = 13. Populations in the southern and western parts of the range are mainly diploid (n = 13) and those in the north and east are mainly tetraploid (n = 26). This species is postulated to be an amphiploid which has resulted from hybridization between the western Cordilleran A. occidentalis (T. & G.) Hook, and the Great Plains species A. falcatus Lindl. Aster ascendens is intermediate between these two species in most morphological traits, but resembles A. occidentalis in ray color and general growth habit, and A. falcatus in the shape of the outer phyllaries. The karyotype of A. ascendens also exhibits a combination of the features found in A. occidentalis and A. falcatus karyotypes. Of the specimens collected for this study, 13% were putative backcrosses of A. ascendens to one of its parent taxa or to a closely related species; this suggests that there is potentially continuing genetic interchange between the species groups that gave rise to A. ascendens.     

Morphological and molecular identification of helminths of the greater bulldog bat Noctilio leporinus (Quiroptera: Noctilionidae) from Campeche,Mexico

Surveys on parasites of bats from the Americas have been conducted, but information on helminths is still scarce, especially in the Neotropical region. In Mexico, there are species of bats that lack of a record for helminth species, such as members of the family Noctilionidae. The present study describes for the first time the helminths of Noctilio leporinus in Campeche, Mexico. In 2017, six specimens of N. leporinus were studied for helminths. The species identification of helminths was based on morphological studies and molecular analysis of fragments of the 28S rDNA. All bat specimens were infected for at least one helminth species. Three helminth taxa were identified: the trematode Pygidiopsis macrostomum, and the nematodes Tricholeiperia cf. proencai, and Heligmonellidae gen. sp. The morphological identification of P. macrostomum was confirmed by sequence analysis of 28S rDNA gene. The phylogeny of P. macrostomum grouped our sequence with other sequences of the same species collected in Brazil. The phylogenetic tree of Heligmonellidae gen. sp. indicated that the helminth belongs to clade formed by the species Odilia bainae, Nippostrongylus magnus and Nippostrongylus brasiliensis of the family Heligmonellidae. The phylogenetic analysis of the 28S sequences of T. cf. proencai did not show any similarity or close affinity with nematodes from which that gene has been sequenced to date. The findings of the present study increase the number of helminth species parasitizing bats in Mexico.     

Comparative study of floral development in Onobrychis melanotricha, Hedysarum varium and Alhagi persarum (Leguminosae: Papilionoideae: Hedysareae)

Floral initiation and development of Hedysarum varium, Onobrychis melanotricha and Alhagi persarum was studied using epi-illumination light-microscopy techniques. The studied species belong to the tribe Hedysareae of the inverted repeat loss clade (IRLC clade), which is characterized by missing the large inverted repeat in the chloroplast genome. The main aim of our study was to determine developmental bases for similarities and differences among the three taxa and to verify the position of Alhagi relative to other genera of the IRLC clade. According to our observations, bracteoles are missing in Onobrychis melanotricha, but are present in the other two species. All three species share unidirectional sepal initiation starting with a median abaxial sepal and bidirectional petal initiation. Stamen initiation is unidirectional in all except in the outer stamen whorl of Hedysarum varium, where it is bidirectional. An important ontogenetic feature in O. melanotricha is the existence of five common primordia, which give rise to petal and stamen primordia. Although in H. varium and O. melanotricha common primordia are observed at some stages in floral organ initiations, in Alhagi all organs are initiated separately. Moreover, overlap in time of floral organs initiation occurs in H. varium and O. melanotricha, but not in A. persarum. The carpel initiates concurrently with the petal primordia in all. It might be presumed that Alhagi is primitive in relation to the other studied Hedysareae taxa, due to the presence of bracteoles, the absence of common primordia, and the lack of overlap in time of different organ initiations.     

Phylogenetic relationships among Indriidae (Primates,Strepsirhini) inferred from highly repeated DNA band patterns

Comparative studies of highly repeated DNA from different species of Indriidae (Primates, Strepsirhini) allowed confirmation of the specific status of Avahi occidentalis, A. laniger and Propithecus tattersalli. The comparison of their band patterns revealed the existence of specific and common bands from which a cladogram of the family is inferred. This cladogram shows that Avahi clade is the sister-group of Indri and Propithecus clade, and that P. verreauxi is related to P. diadema. These results were discussed in view of those obtained from cytogenetic, morphological and molecular data (mitochondrial DNA). This study shows the capacity of the repeated sequence pattern comparison to be used as a tool for confirming taxa status, (taxinomic classification is a primary determinant of management priorities for endangered species, neglect of distinct taxa may lead to their extinction), and for inferring phylogenetic relationships among related species.     

Genetic diversity and relationships among species of the genus Thymus L. (section Serpyllum)

Amplified fragment length polymorphism (AFLP) markers were used to assess genetic relations of 32 Thymus populations belonging to seven species of genus Thymus, section Serpyllum, from Serbia. The collected species belong to three subsections: (1) Alternantes Klokov (Th. pulegioides L.), (2) Isolepides (Borbás) Halácsy (Th. glabrescens Willd., Th. marschallianus Willd., Th. pannonicus All.), and (3) Pseudomarginati (Braun ex Borbás) Jalas (Th. balcanus Borbás, Th. moesiacus Velen., Th. praecox Opiz). Analysis of molecular variance (AMOVA) revealed that most of the genetic diversity was attributable to differences among individuals within populations, while the remaining variability was almost equally distributed among species and among populations within species. The pairwise AMOVA analyses further showed that species differentiation was significant in all cases, except between Th. marschallianus and Th. pannonicus. All the individuals belonging to Th. pulegioides (subsection Alternantes) formed a well-supported clade, the most divergent from all other species in this study. This genetic differentiation of Th. pulegioides is supported by its morphological features because it is the only analyzed taxon having hairs only on the stem edges, and can easily be recognized by this character. According to our results, the subsection Isolepides is polyphyletic. Th. glabrescens is clearly separated from the other two species of this subsection (Th. marschallianus, Th. pannonicus). Although morphologically distinguishable by leaf indumentum, individuals belonging to Th. marschallianus and Th. pannonicus could not be differentiated at the molecular level. The species belonging to subsection Pseudomarginati formed a monophyletic clade, although not supported by high bootstrap value. The clade is further divided into three well-supported clades representing distinct species (Th. balcanus, Th. praecox and Th. moesiacus). In our study, the AFLP markers demonstrated that they could be suitable for studying complex genetic relationships, including frequent interspecies hybridization events, although strict reticulate evolutionary history could not be determined. All the population genetic parameters in the section Serpyllum suggest that the reproductive incompatibility between the species is very weak, which indicates a strong reticulating system, especially in the case when two or more species occur sympatrically.     

三峡库区消落带池杉-土壤碳氮磷生态化学计量特征

为探究三峡库区消落带消落期池杉(Taxodium ascendens Brongn., 1833)及其实生土壤C、N、P生态化学计量特征,于2018年7月对忠县消落带植被修复示范基地3个水淹处理(DS、MS、SS)池杉幼林不同组分(枝条、叶片、根系和土壤)的C、N、P三种元素含量及其化学计量比进行测定分析。结果表明:(1)随着水淹时间和强度的增加,池杉株高、冠幅、基径和胸径均受到一定的抑制,但总体生长良好,与其稳定的化学计量比关系紧密。(2)不同水淹处理组的池杉枝条、叶片中C、N、P含量及其比值分别均无显著性差异(P0.05)。同一水淹处理组的N、P含量表现为叶片根系枝条土壤;除枝条P含量外,其他器官组分C、N、P含量均显著高于土壤组分(P0.05)。(3)池杉各器官N/P比值均远低于临界比率(14),表明池杉的生长可能受N元素限制较为严重。(4)池杉与实生土壤的C、N、P元素内稳性整体表现为PCN,比值内稳性表现为C/NN/PC/P,地上部分(枝条、叶片)C、N、P元素及其比值的稳定性较地下部分(根系)强。(5)冗余分析(RDA)结果表明池杉生态化学计量特征及生长指标与土壤性质密切相关。研究表明,在三峡库区消落带水文多变的环境下,池杉能够有效维持体内化学计量的平衡以响应不同的水淹强度,并且生长良好,是三峡库区消落带植被恢复与重建的优势树种。     

Photobacterium malacitanum sp. nov., and Photobacterium andalusiense sp. nov., two new bacteria isolated from diseased farmed fish in Southern Spain

Three strains, H01100409B^T, H01100413B, and H27100402H^T, were isolated from several internal organs of diseased redbanded seabream (Pagrus auriga) reared in Andalusia (Southern Spain). All strains were studied by phenotypic, including chemotaxonomy, and genomic characteristics. Phylogenetic analysis based on concatenated sequences of six housekeeping genes (gyrB, ftsZ, topA, mreB, gapA, and 16S rRNA) supported the inclusion of the strains within the clade Phosphoreum of the genus Photobacterium, and two of the strains (H27100402H^T and H01100409B^T) formed a tight group separated from the closest species P. aquimaris. Genomic analyses, including average nucleotide identity (ANIb and ANIm) and DNA–DNA hybridization (DDH), clearly separated strains H27100402H^T and H01100409B^T from the other species within the clade Phosphoreum with values below the thresholds for species delineation. The chemotaxonomic features (including FAME analysis and MALDI-TOF-MS) of H27100402H^T and H01100409B^T strains confirmed their differentiation from the related taxa. The results demonstrated that strain H01100413B was classified as P. aquimaris and the strains H27100402H^T and H01100409B^T represented a new species each in the genus Photobacterium, for which we propose the names Photobacterium malacitanum sp. nov., type strain H27100402H^T (=CECT 9190^T = LMG 29992^T), and Photobacterium andalusiense sp. nov., type strain H01100409B^T (=CECT 9192^T = LMG 29994^T).     

NOTES ONXANTHORIATH. FR. III. TWO NEW SPECIES OF THEXANTHORIA CANDELARIAGROUP

Two new species ofXanthoriaare described and illustrated, and comments about their differences from related taxa are provided. These are:X. ascendensfrom bark and shrub remains in sea-shore sand dunes and hillsides in Argentina, Chile and Peru, andX. ucrainica, from bark, stones, and some man-made substrata in the Ukraine, Sweden, United Kingdom, Germany, Austria and Asian Russia.     

Molecular phylogeny of Ceropegia (Asclepiadoideae, Apocynaceae) from Indian Western Ghats

Ceropegia includes more than 200 species distributed in the Old World ranging from the Canary Islands to Australia. In India, there are about 50 species described on a morphological basis as belonging to Ceropegia, and most of them are endemic to the Western Ghats. To investigate evolutionary relationships among Indian Ceropegia taxa and their allies, a phylogenetic analysis was conducted to include 31 Indian taxa of Ceropegia and Brachystelma and their congeners from other geographical regions using nuclear ribosomal internal transcribed spacer (ITS) and three noncoding chloroplast DNA (cpDNA) sequences, including intergenic spacers trnT-L and trnL-F, and trnL intron. The Western Ghats Ceropegia species were found to be most closely related to Indian Brachystelma, with the two genera being placed sister to each other in the ITS phylogeny or with the Brachystelma clade nested within one of the two subclades of Indian Ceropegia in the cpDNA phylogeny. In contrast, Ceropegia species from other regions and African Brachystelma all formed separate clades basal to the Indian Ceropegia–Brachystelma clade. Thus, it can be concluded that the classical morphology-based delineation of the two genera needs revision to reflect their phylogenetic relationships, which are more in accordance with their geographical origin than with morphology.     

Natural hybridization and taxonomy of birches in North Asia

Currently there is a complicated situation in the taxonomy of widespread species such as birch (Betula L.). A large number of species of this genus in Siberia and the Russian Far East has been described, but one encounters an insurmountable obstacle when trying to carry out a more detailed study: the impossibility of finding them for a second time in nature or at least collect plants that have identical characteristics to the typical examples of the described species. All of these species are characterized by a very high variability of traits. These are the traits of Betula pendula and B. microphylla in the southern parts of Central Siberia and Betula pendula (= B. platyphylla) and B. lanata in Eastern Siberia and in the northern regions of the Russian Far East. This suggests intense and widespread hybridization between these species. In connection with this, research of this genus throughout North Asia has begun. These studies have been carried out over 40 years. To date, many works in periodicals and several monographs have been published using the data of numerous expeditions. This paper shows the role of natural hybridization in plant taxonomy using the example of the Siberian species of birch. It is necessary to have a clear understanding of the hybridization processes on large areas that include the ranges of contact species. Solving problems of taxonomy only by studying small local populations is a hopeless pursuit.     

Repeated Reticulate Evolution in North American Papilio machaon Group Swallowtail Butterflies

Hybridization between distinct populations or species is increasingly recognized as an important process for generating biodiversity. However, the interaction between hybridization and speciation is complex, and the diverse evolutionary outcomes of hybridization are difficult to differentiate. Here we characterize potential hybridization in a species group of swallowtail butterflies using microsatellites, DNA sequences, and morphology, and assess whether adaptive introgression or homoploid hybrid speciation was the primary process leading to each putative hybrid lineage. Four geographically separated hybrid populations were identified in the Papilio machaon species group. One distinct mitochondrial DNA clade from P. machaon was fixed in three hybrid taxa (P. brevicauda, P. joanae, and P. m. kahli), while one hybrid swarm (P. zelicaon x machaon) exhibited this hybrid mtDNA clade as well as widespread parental mtDNA haplotypes from both parental species. Microsatellite markers and morphology showed variable admixture and intermediacy, ranging from signatures of prolonged differential introgression from the paternal species (P. polyxenes/P. zelicaon) to current gene flow with both parental species. Divergences of the hybrid lineages dated to early- to mid-Pleistocene, suggesting that repeated glaciations and subsequent range shifts of parental species, particularly P. machaon hudsonianus, facilitated initial hybridization. Although each lineage is distinct, P. joanae is the only taxon with sufficient evidence (ecological separation from parental species) to define it as a homoploid hybrid species. The repetition of hybridization in this group provides a valuable foundation for future research on hybridization, and these results emphasize the potential for hybridization to drive speciation in diverse ways.     

Genetic diversity and specialisation of Eudarluca caricis on some graminaceous Puccinia species

Eudarluca caricis is a common hyperparasite of rusts. A total of 100 cultures were isolated from six Puccinia species or forms growing on 10 species of British grasses at two sites approximately 3 km apart. 82 isolates collected in 2005 were partially sequenced at the ITS locus, and amplified fragment length polymorphism profiles generated for 86 isolates from 2005 and 12 from 2007. Partial ITS sequences of most isolates grouped closely, in a clade with previously reported graminaceous Puccinia isolates and a number of Melampsora isolates. A second clade was very distinct and contained mostly isolates from Puccinia poarum on Poa trivialis. All isolates had distinct AFLP haplotypes. The P. poarum isolates were very distinct from isolates collected from other rusts at the same site. Isolates from P. brachypodii f. sp. arrhenatheri growing on Arrhenatherum elatius in 2005 and 2007 at the same location were distinct (P < 0.001). Isolates from each rust or grass in one year and site were more similar than expected from overall variation between isolates (P < 0.001). Isolates from P. coronata on different grasses clustered together (with isolates from P. brachypodii f. sp. poae-nemoralis), suggesting partial host rust specialisation in E. caricis.     

Molecular phylogenetics of Triculine snails (Gastropoda: Pomatiopsidae) from southern China

Triculine (Gastropoda: Rissooidea: Pomatiopsidae) snails are involved in the transmission of schistosomiasis and paragonimiasis; their distributions are mainly across southeastern Asia and southern China. In the present investigation, partial sequences of COI, 16S, and 28S were examined to infer the phylogenetic relationships among the species rich and poorly understood gastropod. Samples were collected from 12 geographic locations in six provinces of southern China. Several methods such as maximum parsimony, maximum likelihood and distance analysis were used in phylogenetic analyses among these taxa. The resultant phylogenetic trees showed a similar topology irrespective of the phylogenetic methods used. The taxa fell into two clades, with those from Fujian, Guangxi, and Zhejiang Provinces in one clade and those from Hunan, Sichuan and Hubei in the other. Among the taxa in Hubei Province, five formed a monophyletic clade, but Tricula sp. H-SHY fell into a sister clade of Tricula hortensis of Sichuan, whilst Tricula hongshanensis formed a single clade. Sister taxa Tricula pingi and Tricula hsiangi formed well-supported clade within almost all the trees. These results, while preliminary, represent the first attempt to reconstruct a phylogeny for Triculinae across China.     

Early adaptive radiations of Aplodontoidea (Rodentia, Mammalia) on the Holarctic region: systematics, and phylogenetic and paleobiogeographic implications

The Aplodontoidea, now restricted to only one North American species (Aplodontia rufa), have shown a wide Holarctic extension since the Upper Eocene. As their fossil record is poor, their phylogenetic relationships and the origins of their successive radiations remain unclear. We perform here phylogenetic analyses, primarily based on dental evidence (94 dental of 97 characters), restricted to Paleogene and early Miocene taxa (46 taxa) in order to avoid biases introduced by substantially derived (divergent) taxa. We confirm the inclusion of some problematic genera such as Cedromus or Douglassciurus within Sciuroidea rather than in Aplodontoidea. Ephemeromys and Lophallomys appear as the most basal members of the Aplodontoidea, and Epeiromys is the closest outgroup of the Sciuroidea-Aplodontoidea clade. The relationships among the “prosciurines” remain unclear, with paraphyletic genera such as Prosciurus and Haplomys. Their diagnoses are reevaluated and a new genus is described. The Aplodontidae, including the clade of the latter, and Haplomys liolophus display a dichotomy between Ansomyinae and Aplodontinae, the two crown groups. The first clade formed by the European species argoviensis and descendens (referred to a new genus) can be proposed as a sister group of the species of Ansomys. The second branch of the dichotomy includes the European Plesispermophilus and Sciurodon as basal groups. The species of Parallomys do not form a clade, and the genus appears paraphyletic. The last dichotomy separates the Allomys clade from the ‘meniscomyine’ clade. Comparisons of the selected species allow consideration of their patterns of dental evolution (e.g. enlargement of P4, development of a metaloph—protoloph disto-mesial connection, of crescentic shape in main cusps and ectoloph, of a buccal protruding compressed mesostyle, of a metastylid crest or an anterior spur of the hypoconid, etc.). The split between sciuroids and aplodontoids occurred in North America, and then aplodontoids dispersed rapidly throughout the whole Holarctic region. The first aplodontid adaptive radiation took place either in North America or in Asia. Periodic exchanges occurred between Europe, Asia and North America, and the last radiations (meniscomyines) were restricted in North America.     

Phylogeny of Betaphycus (Gigartinales,Rhodophyta) as inferred from COI sequences and morphological observations on B. philippinensis

The phylogeny of the three species that comprise the genus Betaphycus Doty in Silva, Basson and Moe and their phylogenetic relationships with other eucheumatoids are still unresolved. In this study, the utility of the mitochondrial cytochrome oxidase I (COI) marker in resolving their relationships was evaluated. Analyses of the COI sequences from Betaphycus philippinensis Doty and Betaphycus speciosus (Sonder) Doty ex Silva specimens collected from their type localities (Sorsogon, Philippines and Western Australia, respectively) revealed that the two species formed a well-supported clade distinct from Eucheuma J. Agardh and Kappaphycus Doty ex Silva. The genotyped specimens of B. philippinensis were observed to exhibit dorsal protuberances, a characteristic which has been regarded as a key diagnostic feature of Betaphycus gelatinus (Esper) Doty ex Silva. This observation raised the possibility that these two taxa are conspecific. In addition, B. philippinensis specimens were also observed to exhibit morphological features that could be used to distinguish the species from other eucheumatoids, such as the location tetrasporangial nemathecium in the thallus and the presence of apical or lateral pit connections in the tetraspores. The species referred to in the literature as “B. gelatinus” (as Eucheuma gelatinae) collected from northwestern Philippines was identified as a species of Eucheuma based on molecular and morphological evidence. The phylogenetic relationships of this species with other related eucheumatoid taxa were also determined.