A new genus Esexalata gen. n. and two new species (E. beljaevi sp. n., type species, and E. natalyae sp. n.) are described and illustrated from East Malaysia. Males of the new taxa possess unique metathoracic and abdominal androconial structures and wing-like structures on the metascutum. The taxonomic position of the new genus within the family Gelechiidae is discussed.
Matched MeSH terms: Animal Structures/growth & development
The genus Pedionis with its species Pedionis (Pedionis) rufoscutellata Huang & Viraktamath, 1993 is recorded for the first time from Malaysia. A new species Pedionis (Pedionis) spiculata sp. nov. is described and illustrated. An updated species checklist with distribution of the genus Pedionis is also provided.
Matched MeSH terms: Animal Structures/growth & development
All species must partition resources among the processes that underly growth, survival, and reproduction. The resulting demographic trade-offs constrain the range of viable life-history strategies and are hypothesized to promote local coexistence. Tropical forests pose ideal systems to study demographic trade-offs as they have a high diversity of coexisting tree species whose life-history strategies tend to align along two orthogonal axes of variation: a growth-survival trade-off that separates species with fast growth from species with high survival and a stature-recruitment trade-off that separates species that achieve large stature from species with high recruitment. As these trade-offs have typically been explored for trees ≥1 cm dbh, it is unclear how species' growth and survival during earliest seedling stages are related to the trade-offs for trees ≥1 cm dbh. Here, we used principal components and correlation analyses to (1) determine the main demographic trade-offs among seed-to-seedling transition rates and growth and survival rates from the seedling to overstory size classes of 1188 tree species from large-scale forest dynamics plots in Panama, Puerto Rico, Ecuador, Taiwan, and Malaysia and (2) quantify the predictive power of maximum dbh, wood density, seed mass, and specific leaf area for species' position along these demographic trade-off gradients. In four out of five forests, the growth-survival trade-off was the most important demographic trade-off and encompassed growth and survival of both seedlings and trees ≥1 cm dbh. The second most important trade-off separated species with relatively fast growth and high survival at the seedling stage from species with relatively fast growth and high survival ≥1 cm dbh. The relationship between seed-to-seedling transition rates and these two trade-off aces differed between sites. All four traits were significant predictors for species' position along the two trade-off gradients, albeit with varying importance. We concluded that, after accounting for the species' position along the growth-survival trade-off, tree species tend to trade off growth and survival at the seedling with later life stages. This ontogenetic trade-off offers a mechanistic explanation for the stature-recruitment trade-off that constitutes an additional ontogenetic dimension of life-history variation in species-rich ecosystems.
The Pahang Mud Snake (Kualatahan pahangensis) (Tweedie 1946) is a rare species of homalopsid known only from four specimens. In this study we report on a relatively recently collected specimen from Merapoh, Pahang, Peninsular Malaysia, compared with previously collected material. Variations in pholidosis and colour pattern were noted and discussed. The species is morphologically similar in body shape and colour pattern to the sympatric Hypsiscopus plumbeus, for which additional data is presented for Peninsular Malaysian specimens to clarify possible taxonomic confusions. The new record represents a range extension for the species of ~ 47 km to the west and is the third report of the species from the state of Pahang.
Matched MeSH terms: Animal Structures/growth & development
The Austro-Oriental shield-bug genus Axiagastus Dallas, 1851 (Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae: Pentatominae: Axiagastini) is revised and found to be a senior synonym of Indrapura China, 1928, syn. nov. The following Axiagastus species (including five new) are treated: A. cambelli Distant, 1911 (= Pentatoma marmoratum Montrouzier, 1855, syn. nov.), from Australia (Queensland), Papua New Guinea, Solomon Islands, Fiji, Samoa and Vanuatu; A. chandrashekarai Salini, Kment & Roca-Cusachs sp. nov., from Indonesia (Sulawesi); A. dubius Jensen-Haarup, 1937 from ?Singapore; A. klossi (China, 1928, Indrapura), comb. nov., from Indonesia (Sumatra); A. luteipes Salini & Roca-Cusachs sp. nov. from Myanmar; A. mitescens Distant, 1901, from China; A. prathapani Salini & Kment sp. nov., from India (Andaman and Nicobar Islands), Indonesia (Java, Sulawesi, Sumatra), Laos, Malaysia (Malaya, Sabah, Sarawak), and Singapore; A. rosmarus Dallas, 1851 (type species) from the Philippines; A. votypkai Salini & Kment sp. nov., from Indonesia (Papua), Papua New Guinea, Australia (Queensland), and A. yeshwanthi Salini, Kment & Webb sp. nov., from Cambodia, India, Japan (Ryukyus), Taiwan, Thailand and Vietnam. Lectotypes are designated for A. cambelli, A. rosmarus and Pentatoma marmoratum. We maintain the prevailing usage of Axiagastus cambelli and an application to the International Commission on Zoological Nomenclature is prepared to suppress its senior but unused synonym Pentatoma marmoratum. A checklist and key to the included species of Axiagastus are provided. In addition, the species status of Acesines breviceps Stål, 1876 is discussed and the following new subjective synonymy is proposed: Acesines breviceps = Acesines bambusana Distant, 1918, syn. nov. Comments on the collections and type material of William S. Dallas, Francis Walker, Jean X. H. Montrouzier and the Macleay Museum (now part of Chau Chak Wing Museum, University of Sydney) are provided.
Matched MeSH terms: Animal Structures/growth & development
A new species of gekkonid lizard is described from Phnom Khpoh, an isolated karstic hill within an extensive karstic archipelago in Battambang Province, western Cambodia. Phylogenetic analysis using a 1041 base pair fragment of the mitochondrial gene ND2 recovered Hemiphyllodactylus khpoh sp. nov. as the sister species of H. pardalis, found 335 km to the west in southern Thailand. Analyses of meristic, morphometric, and categorical characters of morphology and color pattern differentiated H. khpoh sp. nov. from both the morphologically distinct H. pardalis and the very similar Hemiphyllodactylus sp. from Phnom Kulen, approximately 130 km to the northeast in Siem Reap Province. The statistical morphological analyses and comparisons also showed that H. khpoh sp. nov. is well-differentiated from the recently described H. bokor and H. samkos from the nearby Cardamom Mountains as well from species in clade 6-a recently designated lineage from China, Laos, and Vietnam to which H. khpoh sp. nov. + H. pardalis compose the sister lineage. The discovery of H. khpoh sp. nov. highlights the rich but understudied biodiversity of karstic landscapes, emphasizing the need for continued research in the extensive unexplored karstic archipelago of western Cambodia. This region is poised to reveal unparalleled gekkonid diversity, comparable to that observed in the similarly sized karstic archipelago of the Salween Basin in southern Myanmar. Although karstic landscapes are proving to be some of the most biodiverse habitats on the planet, less than 1% of them are afforded any formal protection.
Matched MeSH terms: Animal Structures/growth & development
Colonisation of maxillofacial prostheses by Candida albicans is conventionally attributed to surface roughness, porosity, and wettability properties that explain where cells accumulate rather than why particular surfaces promote biofilm formation. This review reinterprets the in vitro literature on C. albicans adhesion to maxillofacial silicone elastomers through the underlying molecular events. Nine eligible studies identified through searches of PubMed, Scopus, and ScienceDirect conducted on 31 March 2026 were appraised with the QUIN tool and mapped onto a six-stage molecular cascade: surface conditioning, reversible attachment, adhesin engagement, morphogenesis, matrix assembly, and antifungal tolerance. Three conclusions emerged. First, C. albicans senses topography through mechanically gated calcium channels and converts mechanical contact into hyphal commitment; roughness is therefore a morphogenetic cue that surface finishing can withhold. Second, the tested agents-silver nanoparticles, chitosan, thymol, and carvacrol, and a Lactobacillus brevis-derived biosurfactant-engage distinct molecular targets and are not mechanistically interchangeable. Third, most included studies evaluated silicone without salivary conditioning, whereas the clinical interface is an acquired salivary pellicle that supplies both adhesion receptors and antimicrobial peptides. All studies fell within the medium risk-of-bias band (50.0-62.5%). Pellicle-conditioned models and molecular readouts are required. The mechanistic interpretations discussed in this review are derived from the broader C. albicans literature and have not been directly demonstrated on maxillofacial silicone elastomers.
Sphenomorphus stellatus was once thought to be a wide-ranging species distributed across many parts of Indochina, the Thai-Malay Peninsula and Borneo. However, recent integrative taxonomic studies on the species have revealed its underestimated diversity and new species previously masquerading under this nomen in mainland Southeast Asia. This prompted the re-examination of a specimen identified as S. stellatus from Borneo, and morphological analyses revealed that it is not conspecific with topotypic specimens of the species from Peninsular Malaysia or other congeners of the complex, and is therefore described herein as new. Sphenomorphus striatafaucium sp. nov. can be differentiated from its congeners by a combination of known adult SVL of 57.4 mm; frontal scale undivided; prefrontals in broad contact; four supraoculars; two loreals; eight superciliaries; no enlarged, clear, central window in lower eyelid; seven or eight supralabials; six infralabials; three pairs of chinshields; 24 midbody scale rows; 62 paravertebrals, not wider than other dorsals; 66 ventrals; two enlarged precloacals; 19 subdigital lamellae on fourth toe; and bold and distinct striped pattern on the chin and throat. The discovery of S. striatafaucium sp. nov. increases the number of endemic Sphenomorphus skinks in Borneo to 13 species. The discovery underscores the underestimated diversity of reptile and amphibian species in Borneo, and the need for more concerted efforts to conduct fieldwork and systematic studies to not only improve the understanding of their relationships and ecologies, but also better protect their habitats that are increasingly threatened by anthropogenic pressures. This study also highlights the value of well-curated natural history collections in museums that still harbour new species awaiting discovery.
Matched MeSH terms: Animal Structures/growth & development
The Hemipeplinae (Coleoptera: Mycteridae) fauna of the world was reviewed, based on external morphology, morphometrics of head, pronotum, and elytra, and examination of male genitalia of adults. One new genus and twenty-nine new species are described (country of type locality given in parentheses). From the New World, five new species are described: Hemipeplus anishae KC & Pollock, sp. nov. (Belize), H. belizensis KC & Pollock, sp. nov. (Belize), H. brasiliensis KC & Pollock, sp. nov. (Brazil), H. pollocki KC, sp. nov. (Mexico), and H. pseudoquadricollis KC & Pollock, sp. nov. (Brazil). From the Afrotropics, one monotypic genus with its species is described: Eurypeplus quadrimaculatus KC & Pollock, gen. nov., sp. nov. (Madagascar). Twenty-three new species are described from the Austro-Oriental region: H. antennatus KC & Pollock, sp. nov. (Solomon Islands), Hemipeplus bromineus KC & Pollock, sp. nov. (Indonesia: Papua), H. bucculentus KC & Pollock, sp. nov. (Vietnam), H. craigi KC & Pollock, sp. nov. (Papua New Guinea), H. enarotaliensis KC & Pollock, sp. nov. (Indonesia: Papua), H. exhaustus KC & Pollock, sp. nov. (Papua New Guinea), H. gressitti KC & Pollock, sp. nov. (Indonesia: Papua), H. harkoneni KC & Pollock, sp. nov. (Philippines), H. heisenbergi KC & Pollock, sp. nov. (Papua New Guinea), H. joerebeccae KC & Pollock, sp. nov. (Philippines), H. labuanensis KC & Pollock, sp. nov. (Malaysia), H. manusicus KC & Pollock, sp. nov. (Papua New Guinea), H. monteithi KC & Pollock, sp. nov. (Australia), H. neoaustralicus KC & Pollock, sp. nov. (Australia), H. neoguineensis KC & Pollock, sp. nov. (Papua New Guinea), H. nigerrimus KC & Pollock, sp. nov. (Philippines), H. palawanensis KC & Pollock, sp. nov. (Philippines), H. philippinensis KC & Pollock, sp. nov. (Philippines), H. pseudoklematanicus KC & Pollock, sp. nov. (Malaysia), H. rugosus KC & Pollock, sp. nov. (Malaysia), H. luzonensis KC & Pollock, sp. nov. (Philippines), H. saymyname KC & Pollock, sp. nov. (Malaysia), and H. sinensis KC & Pollock, sp. nov. (China). Range extensions of several previously described species are documented: Hemipeplus thomasi Pollock, 1999 is recorded from Texas (Cameron Co.) previously known only from Tamaulipas, Mexico; H. bolivianus Pollock, 1999 is newly recorded from Ecuador; H. alluaudi Grouvelle, 1923 is recorded from Malawi, previously known from southern Kenya; H. miyamotoi Kamiya, 1961 is recorded from Taiwan, previously known from Japan; H. klematanicus (Gestro, 1873), previously known only from Malaysia (Sarawak) is recorded from Thailand and Indonesia. Hemipeplus marginipennis (LeConte, 1856) is proposed as a junior synonym of H. hemipterus Lacordaire, 1854, syn. nov.; a neotype is designated for the latter. Lectotypes for H. australicus Arrow, 1930 and H. nuciferae Arrow, 1930 are designated. To supplement the descriptions of new taxa, images of different aspects of their habiti and male genitalia (whenever possible) are provided along with taxonomic keys for all the known taxa of Hemipeplinae.
Matched MeSH terms: Animal Structures/growth & development
Root-specific or preferential promoters are essential to genetically modify plants with beneficial root traits. We have characterised the promoter from an oil palm metallothionein gene (EgMT) and performed a serial 5' deletion analysis to identify the region(s) essential for transgenes expression in roots. Stable functional characterisation of tobacco transgenic lines using the T1 generation showed that a deletion construct, designated as RSP-2D (1107 bp), directed strong GUS expression at all stages of root development, particularly in mature roots. Other constructs, RSP-2A (2481 bp) and RSP-2C (1639 bp), drove GUS expression in roots with an intensity lower than RSP-2D. The promoter activity was also detectable in seed pods and immature seeds, albeit at lower levels than CaMV35S. The promoter activity may also be induced by wounding as intact GUS staining was observed at the flower- and leaf-cutting sites of T1 samples carrying either RSP-2C or RSP-2D constructs. The promoter sequence contains cis-acting elements that may act as negative regulators and be responsible for root specificity. The results further indicated that the 5' UTR and ATATT sequences are essential for strong promoter activity. This study highlights the potential of RSP-2D promoter as a tool for modifying root traits through genetic engineering.
Matched MeSH terms: Arecaceae/growth & development
Banana is a premier fruit crop in many parts of the world especially Southeast Asia. The demand for banana has contributed to significant national income to primary banana producers in the SEA region such as the Philippines, Indonesia, Thailand, Vietnam, and Malaysia. However, the widely traded banana industry is plagued by numerous threats including pests and diseases, post-harvest issues and extreme climate vulnerability. To address these challenges, new breeding techniques such as gene editing have been explored for breeding programs to develop improved banana varieties. The first gene-edited non-browning banana has been deregulated in the Philippines recently, and more regulatory applications are expected to submit for approvals soon. Hence, it is timely to review the policy options for gene editing that have been adopted and discussed in the Southeast Asian countries and highlight the implications of differing regulatory approaches to gene editing for trading activities. Positive stakeholders' perceptions and public acceptance are key factors in allowing the benefits of gene editing and thus appropriate outreach strategies are important to gain acceptance and avoid the "GMO stigma" that may be associated with gene-edited products.
Matched MeSH terms: Plants, Genetically Modified/growth & development
Seeds are essential for plant reproduction, ensuring species survival and dispersal while adapting to diverse environments throughout a plant's life. Proteomics has emerged as a powerful tool for deciphering the complexities of seed growth, germination, and stress responses. Advanced proteomic technologies enable the analysis of protein changes during germination, dormancy, and ageing, enhancing our understanding of seed lifespan and vitality. Recent studies have revealed detailed insights into metabolic processes and storage protein profiles across various plant species. This knowledge is crucial for improving seed storage, conserving quality, and maintaining viability. Additionally, it contributes to sustainable agriculture by identifying stress-responsive proteins and signalling pathways that can mitigate stress and enhance farming practices. This review highlights significant advancements in seed proteomics over the past decade, discussing critical discoveries related to storage proteins, protein interactions, and proteome modifications due to stress. It illustrates how these insights transform seed biology, boosting productivity, food security, and environmentally friendly practices. The review also identifies existing knowledge gaps and provides direction for future research, underscoring the need for continued interdisciplinary collaboration in this dynamic field.
Matched MeSH terms: Crops, Agricultural/growth & development
The tropical forests of Borneo and Amazonia may each contain more tree species diversity in half a square kilometre than do all the temperate forests of Europe, North America, and Asia combined. Biologists have long been fascinated by this disparity, using it to investigate potential drivers of biodiversity. Latitudinal variation in many of these drivers is expected to create geographic differences in ecological and evolutionary processes, and evidence increasingly shows that tropical ecosystems have higher rates of diversification, clade origination, and clade dispersal. However, there is currently no evidence to link gradients in ecological processes within communities at a local scale directly to the geographic gradient in biodiversity. Here, we show geographic variation in the storage effect, an ecological mechanism that reduces the potential for competitive exclusion more strongly in the tropics than it does in temperate and boreal zones, decreasing the ratio of interspecific-to-intraspecific competition by 0.25% for each degree of latitude that an ecosystem is located closer to the Equator. Additionally, we find evidence that latitudinal variation in climate underpins these differences; longer growing seasons in the tropics reduce constraints on the seasonal timing of reproduction, permitting lower recruitment synchrony between species and thereby enhancing niche partitioning through the storage effect. Our results demonstrate that the strength of the storage effect, and therefore its impact on diversity within communities, varies latitudinally in association with climate. This finding highlights the importance of biotic interactions in shaping geographic diversity patterns, and emphasizes the need to understand the mechanisms underpinning ecological processes in greater detail than has previously been appreciated.
The recurrent patterns in the commonness and rarity of species in ecological communities--the relative species abundance--have puzzled ecologists for more than half a century. Here we show that the framework of the current neutral theory in ecology can easily be generalized to incorporate symmetric density dependence. We can calculate precisely the strength of the rare-species advantage that is needed to explain a given RSA distribution. Previously, we demonstrated that a mechanism of dispersal limitation also fits RSA data well. Here we compare fits of the dispersal and density-dependence mechanisms for empirical RSA data on tree species in six New and Old World tropical forests and show that both mechanisms offer sufficient and independent explanations. We suggest that RSA data cannot by themselves be used to discriminate among these explanations of RSA patterns--empirical studies will be required to determine whether RSA patterns are due to one or the other mechanism, or to some combination of both.
Old-growth tropical forests are widely recognized as being immensely important for their biodiversity and high biomass1. Conversely, logged tropical forests are usually characterized as degraded ecosystems2. However, whether logging results in a degradation in ecosystem functions is less clear: shifts in the strength and resilience of key ecosystem processes in large suites of species have rarely been assessed in an ecologically integrated and quantitative framework. Here we adopt an ecosystem energetics lens to gain new insight into the impacts of tropical forest disturbance on a key integrative aspect of ecological function: food pathways and community structure of birds and mammals. We focus on a gradient spanning old-growth and logged forests and oil palm plantations in Borneo. In logged forest there is a 2.5-fold increase in total resource consumption by both birds and mammals compared to that in old-growth forests, probably driven by greater resource accessibility and vegetation palatability. Most principal energetic pathways maintain high species diversity and redundancy, implying maintained resilience. Conversion of logged forest into oil palm plantation results in the collapse of most energetic pathways. Far from being degraded ecosystems, even heavily logged forests can be vibrant and diverse ecosystems with enhanced levels of ecological function.
Bactrocera carambolae is a highly polyphagous fruit pest of agricultural importance. This study reports the bacterial communities associated with the developmental stages of B. carambolae. The microbiota of the developmental stages were investigated by targeted 16S rRNA gene (V3-V4 region) sequencing using the Illumina MiSeq. At 97% similarity, there were 19 bacterial phyla and unassigned bacteria, comprising 39 classes, 86 orders, 159 families and 311 genera. The bacterial composition varied among the specimens of developmental stage and across developmental stages as well as exuviae. Four phyla of bacteria (with relative abundance of ≥1% in at least one specimen)-Actinobacteria, Bacteroidetes, Firmicutes and Proteobacteria-were recovered from the larva, pupa, adult stages and exuviae. Proteobacteria was the predominant phylum in all the developmental stages as well as the exuviae. Enterobacteriaceae (Proteobacteria) was the predominant family in the adult flies while the family [Weeksellaceae] (Bacteroidetes) was predominant in the larval and pupal stages. Among the genera occurring in more than one developmental stage of B. carambolae, Erwinia was more abundant in the larval stage, Halomonas more abundant in adult female, Stenotrophomonas more abundant in adult male, and Chryseobacterium more abundant in the larval and pupal stages. The results indicate transmission of bacteria OTUs from immatures to the newly emerged adults, and from exuviae to the environment.
A new black fly species, Simulium haiduanense Takaoka, Low & Huang (Diptera: Simuliidae), is described on the basis of females, males, pupae, and mature larvae from Taiwan. This new species is placed in the Simulium argentipes species-group of the subgenus Simulium (Diptera: Simuliidae) and is characterized by the yellowish female legs, ovipositor valves rounded apically and with its inner margin concave, claw with a small subbasal tooth, male style without a basal protuberance, pupal gill with eight filaments, corbicular cocoon, and larval abdomen lacking paired protuberances. It represents the first record of the S. argentipes species-group from Taiwan. Taxonomic notes are given to separate this new species from all eight species in the same species-group. The phylogenetic relationships of this new species with four related species are presented.
KEY MESSAGE: Morphological and transcriptomic evidences provide us strong support for the function of ANAC019 in reproductive development under drought stress. Plants are sensitive to drought conditions, particularly at the reproductive stage. Several studies have reported drought effects on crop reproductive development, but the molecular mechanism underlying drought response during reproduction is still unclear. A recent study showed that drought induces in Arabidopsis inflorescence increased expression of many genes, including ANAC019. However, the function of ANAC019 in drought response during reproductive development has not been characterized. Here, we report an investigation of the ANAC019 function in the response to drought during reproduction. ANAC019 is preferentially expressed in the inflorescence compared with the leaf, suggesting possible roles in regulating both stress response and flower development. The anac019 mutant was more sensitive to drought than WT plant, and exhibited a delay in recovery of floral organ development under prolonged drought stress. Moreover, many fewer genes were differentially expressed in the anac019 inflorescence under drought than that of WT, suggesting that the mutant was impaired in drought-induced gene expression. The genes affected by ANAC019 were associated with stress and hormone responses as well as floral development. In particular, the expression levels of several key drought-induced genes, DREB2A, DREB2B, ARF2, MYB21 and MYB24, were dramatically reduced in the absence of ANAC019, suggesting that ANAC019 is an upstream regulator these genes for drought response and flower development. These results provide strong support for the potential function of ANAC019 in reproductive development under drought stress.
BACKGROUND: Methicillin-resistant Staphylococcus aureus (MRSA) biofilm producers represent an important etiological agent of many chronic human infections. Antibiotics and host immune responses are largely ineffective against bacteria within biofilms. Alternative actions and novel antimicrobials should be considered. In this context, the use of phages to destroy MRSA biofilms presents an innovative alternative mechanism.
RESULTS: Twenty-five MRSA biofilm producers were used as substrates to isolate MRSA-specific phages. Despite the difficulties in obtaining an isolate of this phage, two phages (UPMK_1 and UPMK_2) were isolated. Both phages varied in their ability to produce halos around their plaques, host infectivity, one-step growth curves, and electron microscopy features. Furthermore, both phages demonstrated antagonistic infectivity on planktonic cultures. This was validated in an in vitro static biofilm assay (in microtiter-plates), followed by the visualization of the biofilm architecture in situ via confocal laser scanning microscopy before and after phage infection, and further supported by phages genome analysis. The UPMK_1 genome comprised 152,788 bp coding for 155 putative open reading frames (ORFs), and its genome characteristics were between the Myoviridae and Siphoviridae family, though the morphological features confined it more to the Siphoviridae family. The UPMK_2 has 40,955 bp with 62 putative ORFs; morphologically, it presented the features of the Podoviridae though its genome did not show similarity with any of the S. aureus in the Podoviridae family. Both phages possess lytic enzymes that were associated with a high ability to degrade biofilms as shown in the microtiter plate and CLSM analyses.
CONCLUSIONS: The present work addressed the possibility of using phages as potential biocontrol agents for biofilm-producing MRSA.
We screened more than 60 Malaysian plants against two species of insects and found that Melicope subunifoliolata (Stapf) T.G. Hartley (Rutaceae) showed strong feeding deterrent activity against Sitophilus zeamais Motsch. (Curculionidae) and very good larvicidal activity against Aedes aegypti L. (Diptera). One anti-insect compound, meliternatin (3,5-dimethoxy-3',4',6,7-bismethylendioxyflavone) (6) and six other minor polyoxygenated flavones were isolated from M. subunifoliolata.