Journal of Threatened Taxa | www.threatenedtaxa.org | 26 April 2026 | 18(4): 28696–28702

 

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) 

https://doi.org/10.11609/jott.9997.18.4.28696-28702

#9997 | Received 02 July 2025 | Final received 21 March 2026| Finally accepted 09 April 2026

 

 

Unveiling genital specializations in Megascolex travancorensis (Oligochaeta: Megascolecidae) through scanning electron microscopy

 

Jaya Manazhy 1, Sona Sajeev 2, Aja Manazhy 3 , John Warren Reynolds 4  & Santhosh Punnakattu Parambil 5          

 

1,2 Department of Zoology, Sree Kerala Varma College, Thrissur, University of Calicut, Kerala 680011, India.

3,5 Department of Zoology, Sree Krishna College, Guruvayur, University of Calicut, Kerala 680602, India.   

4 Oligochaetology Laboratory, 9-1250 Weber Street East, Kitchener, ON N2A 4E1, Canada.

4 New Brunswick Museum, 277 Douglas Street, Saint John, NB E2K lE5, Canada.

1 jayamanazhy@gmail.com (corresponding  author), 2 sonasajeev72@gmail.com, 3 ajamanazhy@gmail.com, 4 john.w.reynolds1941@gmail.com, 5 santhuptb@gmail.com

 

 

Editor: Ankit Kumar Singh, Tribhuvan University, Kathmandu, Nepal.     Date of publication: 26 April 2026 (online & print)

 

Citation: Manazhy, J., S. Sajeev, A. Manazhy, J.W. Reynolds & S.P. Parambil (2026). Unveiling genital specializations in Megascolex travancorensis (Oligochaeta: Megascolecidae) through scanning electron microscopy. Journal of Threatened Taxa 18(4): 28696–28702. https://doi.org/10.11609/jott.9997.18.4.28696-28702

  

Copyright: © Manazhy et al. 2026. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by providing adequate credit to the author(s) and the source of publication.

 

Funding: The authors declare that no funding was received for this study.

 

Competing interests: The authors declare no competing interests.

 

Author details: Jaya Manazhy is an Assistant Professor in the Department of Zoology at Sree Kerala Varma College, Thrissur, Kerala and Aja Manazhy is an Assistant Professor (on Contract) in the Department of Zoology at Sree Krishna College, Guruvayur, Kerala. Both are academicians actively engaged in research, with a primary focus on animal systematics and biology. Sona Sajeev is an environmental educator with a primary focus on wildlife studies and forest exploration. John Warren Reynolds Ph. D, LL. B is the director of Oligochaetology Laboratory, Kitchener and Research Associate at the New Brunswick, Saint John, Canada, whose speciality is the taxonomy and biogeography of the Oligochaeta. Dr. Santhosh P.P. is an Associate Professor and head of the Department of Zoology, Sree Krishna College, Guruvayur, serving as Head since 2019. His research interests include General Zoology, Acarology, and Entomology.

 

Author contribution: Jaya Manazhy and Aja Manazhy conceptualized the study and contributed to the review and editing of the manuscript. Sona Sajeev conducted the field study, performed SEM analysis, curated the data, prepared visualizations, and wrote the original draft. John Warren Reynolds provided resources and contributed to the review and editing. Santhosh Punnakattu Parambil contributed to data analysis and manuscript editing

 

Acknowledgments: We express our sincere gratitude to Sree Kerala Varma College, University of Calicut, Kerala for providing the necessary facilities, and to CLIF, University of Kerala, Kerala for facilitating high-resolution SEM imaging.

 

 

Abstract: When an earthworm reaches reproductive age, its clitellum, also referred to as the genital collar, observably enlarges and becomes more glandular. Genital features in Megascolecidae, including male and female pores, papillae, grooves, and glandular thickenings, show species-specific patterns linked to reproduction, thereby serving as important characters in taxonomy and systematics. This work uses comprehensive scanning electron microscopy (SEM) images to investigate the male reproductive anatomy of Megascolex travancorensis Michaelsen, 1910. A slit-shaped male pore located on a raised genital papilla, paired prostatic ducts, a seminal groove, retracted penial setae, and eye-like genital marks on segment 18 are among the distinctive morphological features highlighted in the investigation. Fine-scale structural adaptations have been shown by SEM, which has also helped to classify accurately the genus Megascolex taxonomically and provide important information about reproductive morphology.

 

Keywords: Clitellum, earthworms, genital papilla, penial setae, Prostatic ducts, reproduction, Seminal groove, taxonomy, Tubercula Pubertatis.

 

 

 

INTRODUCTION

 

Hermaphroditism and the presence of a clitellum, a specialized reproductive structure, are the main characteristics that set oligochaetes apart from polychaetes (Reynolds 1974). With more than 5,000 species of terrestrial earthworms, the oligochaetes make up about one-third of the phylum and are the second largest group within Annelida (Reynolds & Wetzel 2025). Their wide range of terrestrial environments highlights their astounding adaptability and ecological significance. The clitellum, a noticeable glandular growth near the front end of their reproductive system, is a crucial component that emerges at sexual maturity. Earthworms produce eggs for a large portion of the year and are regarded as either continuous or semi-continuous breeders (Olive & Clark 1978). Their reproductive activity is greatly influenced by environmental conditions, especially soil temperature and moisture content (Reynolds 2022).

The ventral or ventrolateral surface contains pairs of genital markings, which are external apertures for reproductive exchange, indicating their hermaphrodite characteristics. These apertures and related features display morphological characteristics unique to each species, which are useful in systematics and taxonomy. Genital features that are frequently connected to glands, such as discs, pits, grooves, or pads, are useful for identifying species (Blakemore 2006). Male genital area configuration and sperm transfer requirements are frequently correlated with the shape and location of these characteristics. Male pore arrangements vary; the megascolecine type has both male and prostatic pores on segment 18, but the acanthodriline, microscolecine, and balantine forms have them separated or relocated to segments 17 or 19, occasionally connected by seminal grooves or changed by segmental displacement. In Megascolecidae, the male pores may be associated with prostatic pores, both of them may appear on raised papillae or ridges, and posing tubular or racemose prostates which in turn associated with the posterior ends of vasa defentia. The megascolecids often possess various markings at sexual maturity in the form of tubercles which are the glandular thickenings on the ventral surface and situated both sides or on the clitellum (Figure 1).

There are studies available to identify oligochaetes morphologically across various regions of the world (Gates &  Reynolds 2017; Khan et al. 2022; Kamal et al. 2023; Rathour & Julka 2024; Narayanan et al. 2025). There is still a dearth of ultrastructural research on earthworm reproductive architecture in the body of literature, which indicates a lack of thorough microscopic morphological characterization. In-depth ultrastructural examination of genital characteristics, especially with scanning electron microscopy (SEM), allows for accurate characterization of minute surface details such glandular areas, pore shape, and related textures, these are the aspects that conventional light microscopy frequently fails to discern. Proper sample preparation is essential to ensure accurate SEM imaging, minimizing artifacts and enhancing structural clarity. SEM is, therefore, an essential tool for researching the reproductive morphology of earthworms and enhancing species-level identification. The present study aims to characterize the ultrastructural morphology of the selected species, thereby providing insights into their taxonomic and functional significance.

 

 

Materials and Methods

 

Earthworms with well-developed clitellum were obtained from the soil by digging and hand sorting methods (Reynolds 1977, 2022), and also searching in organic micro habitats like leaf litter. Out of the 30 samples collected, ten well-developed clitellate adult specimens were preserved in 10% formaldehyde for further identification using a light microscope and each sample was assigned a code . The type specimens were deposited in Zoology Museum, Sree Kerala Varma College, Thrissur, Kerala with deposition number THS201. For SEM analysis, two specimens were fixed in 5% glutaraldehyde (C5H8O2) for 30 minutes to preserve ultrastructural details. Following fixation, the dissected clitellum was washed thoroughly and dehydrated using an ascending ethanol series. Specimens are mounted on to SEM stubs using conductive adhesive like carbon taps and coated with a thin layer of conductive material (gold/palladium) using a sputter coater to enhance image quality and reduce charging effects. Prepared samples are placed in the SEM chamber, where they are scanned by the electron beam to generate high-resolution images. The secondary soil data from various localities were obtained from the data provided by the Soil Survey Organisation Agriculture Department (Premachandran 2007). Observations were carried using Carl Zeiss EVO 18 Research SEM (Central laboratory for Instrumentation and facilitation, University of Kerala) and magnified the sample from 10X to 5K X.

 

 

RESULTS

 

Morphological identification by Light microscopy

 

Family Megascolecidae Rosa, 1891

(Diagnostic features from: descriptions of Stephenson 1923; Gates 1972; Blakemore 2006; Reynolds 2022 and also present observation)

 

Diagnostic Features

Dorsal pores: present

Male pores: male pores may be united with prostatic pores, paired or unpaired, commonly on 18, rarely on 19 or 20, behind female pores.

Setae: lumbricine or perichaetine.

Clitellum: annular, multilayered.

Nephridia: holoic or meroic.

Intestinal origin: behind ovarian segment.

Gizzard: Oesophageal gizzard(s) present; intestinal gizzard(s): present, or absent.

Last hearts: behind segment 11.

Calciferous glands: present or absent.

Ovaries: in 13, mostly fan shaped and with numerous egg strings.

Spermathecae: single, paired, or multiple; diverticulate (or rarely with intramural sperm chambers),

Prostates: present, tubular to racemose.

Remarks: The position of male pore and nature of prostate are the diagnostic features.

 

Genus Megascolex Templeton, 1844

(Diagnostic Features from: descriptions of Stephenson 1923; Aiyer 1929 and also present observation)

1844.   Megascolex –Templeton, Proc. Zool. Soc. London, 12(1844): 89.

1895.     Megascolex – Beddard, Monogr. Oligochaeta, p. 370.

1900.     Megascolex – Michaelsen, Oligochaeta, Das Tierrich, p. 212.

1907.     Megascolex – Michaelsen, Fauna S W Australia 1: 168.

1909.     Megascolex + Lampito – Michaelsen, Mem. Indian Mus. 1: 178.

1916.     Megascolex – Michaelsen, Mjobeg s Australia Exp. 52: 57.

2009.  Megascolex – Ramasamy, Doctor of Philosophy in Zoology, Tamilnadu, India, p. 99.

 

Diagnostic features

Setae: perichaetine.

Spermathecal pores: 1–5 pairs between segments 4–9.

Gizzard: one in 5, 6 or 7.

Nephridia: meroic.

Prostate: racemose to tubulo-racemose.

Previous records: India (Southern India, Bombay, eastern Himalaya), Australia, Ceylon, Tasmania, northern Island of New Zealand, southwestern Asia, and the Malay Archipelago.

 

 Megascolex travancorensis Michaelsen, 1910

 Redescription

1910.    Megascolex travancorensis f. typica Michaelsen, Abh. Ver. Hamburg 19 (5): 72.

1913.    Megascolex travancorensis var. typicus- Michaelsen, Mt. Mus. Hamburg 30(2): 85.

Collectors’ name: Sona Sajeev.

Date of collection: 29.iv.2024.

General habitat: coconut plantation (Cocos sp.), cultivated area.

Soil texture: gravelly sandy clay, sandy clay loam, clay loam, gravelly clay and sand.

Physiography: low land, mid upland, mid land, mountainous region (classification based on Soil Survey Organisation Agricultural Department, Premachandran 2007).

(Diagnostic Features from: descriptions of Stephenson 1923 and also present observation)

 

External Features

Size: 80–140 mm; preclitellar region: 2 mm; clitellar region: 3 mm; post clitellar region: (1–2 mm).

Number of segments: 120–280, segment 2–5 biannular; 5–13 triannular.

Pigmentation: dark grey before preservation, unpigmented (Image 1).

Prostomium: epi-tanylobous.

Setae: perichaetine, 16 setae per segment in the middle of the body, However, anterior segments (= up to the segment 25) the setae in lumbricine arrangement (Figure 2).

First dorsal pore: 4/5 (3/4 in one specimen).

Clitellum: well, developed in segments 13–18 (= 6); in some specimens overlapping the male genital markings.

Male pores: in 18 on slightly elevated cushions extended to segment 19, in ab, bc and extends to ¾ of cd on each side which are ellipsoidal, their inner borders approximated and parallel, both cushions together almost fill up a somewhat depressed median area, which is bounded laterally and in front by a slight wall.

Female pores: paired.

Spermathecal pores: two pairs, between a and b, in 7/8 and 8/9, about ½ of the circumference apart.

Genital marking: pair of elevated markings in segments 17, 18 and 19 at more or less at cd in which it is highly ornamented with folding and wrinkles forming the shape of an eye at the base (= 19).

 

Internal Anatomy

 Septa: 6/7 – 12/13 highly thickened, 13/14 and 14/15 moderately thickened.

 Gizzard: large in 6.

 Calciferous glands: absent.

 Testis and funnels: free in 10 and 11.

 Seminal vesicle: compactly racemose in 11 and 12.

 Spermathecae: spermathecal ampulla large, pear-shaped and usually much bent at its ectal end; duct still thinner, very short, mostly concealed in the body-wall; diverticulum enters the ectal end of ampulla; is narrowly club-shaped and somewhat bent at its ectal end; a mass at the ectal end of ampulla seems to represent an incompletely formed spermatophore.

 Prostates: racemose in the 17 and part of the following segments with three to five incisions to form lobes.

 Ovary: two pairs in 13 and 14.

 Penial setae: absent.

Remarks: The spermathecae similar to Megascolex konkanensis. The form of tubercula pubertatis is also a notable feature of the type species M. travancorensis. In some specimens, we noticed a digitiform projection in the segments 15/16/17/18. The actual function of these structures is not known and may be a genital marking or an artifact.

Previous records: Southern India.

Present collection: Thiruvananthapuram, Kollam, Pathanamthitta (south), Thrissur, Palakad (central), Kerala, southern India.

 

Ultramorphological analysis of genital pore using SEM

Male genital area is highly ornamented with folding, wrinkles, and grooves associated with more pronounced genital marking (Image 2). Although the eye shaped marking is visible by light microscopic analysis, the associated pores and furrows are inconspicuous. The exact segmental location, shape, pattern, and associated structures furnish useful taxonomic characteristics. The extension of clitellar glandular tissue in segments 13–18 overlapping the male genital markings on segment 18 suggests a close structural association between the clitellum and reproductive pores reflecting the genital specialization especially in the genus Megascolex. By ultrastructural analysis, the male pore is clearly visible on segment 18 as a small slit with a magnification of 60 X and slightly raised on a male genital papilla that slightly extended to segment 19 (Image 3A,B). Here, tubercula pubertatis appears as a pair of glandular thickened ridges arises on the segments 17–19. Seminal groove seen as a shallow channel that starts from the male pore and runs anteriorly towards the clitellum (Image 3B, 370 X) and that helps in transferring sperm during copulation. Prostatic ducts appear as two lateral grooves one above the other from the paired prostatic lobes (Image 4). A deep pore is seen near to the prostatic pore may be the indication of the retracted penial setae (Image 4, 5KX).

 

DISCUSSION            

 

Similar possibility discussed by Gates (1972) in ‘Burmese earthworms’ that male chambers may be eversible into characteristic porophores and may contain protrusible penes. Erséus (1981) discussed that Bathydrilus rohdei is provided with an X-shaped mid-ventral slit that, corresponds to the location of the tips of the penial setae of the mate. There are previous reports available for specialized genital setae and related glands that make up the sexual papillae may be used to inject sperms into the mate (Cosín et al. 2011). Ambreen et al. (2004) reported that the variations in genital papilla in the genus Pheretima (sensu lato) may contribute to species identification. Similar to our study, Yáñez et al. (2006) and Caramelo & Martínez-Ansemil (2012) used scanning electron microscopy to study particular mating structures and sperm transmission processes in aquatic microdriles, exposing copulation-related ultrastructural adaptations and specialized attachment features.

It should be noted that the families Ocnerodrilidae, Acanthodilidae, Octochaetidae, and Megascolecidae, the male pores are associated with the prostatic pores (Julka 1988). In megascolecids with the ‘megascolecine’ arrangement showed each type of gland connects to the male pores by a muscular prostatic duct that receives sperm from the testis via the vasa deferentia that often joins the duct near the junction with the gland and one pair of male and prostatic pores combined on segment 18 (Blakemore 2006). Here we can observe that, although M. travancorensis, there are separate pores of prostatic ducts and later joins with vasa deferentia to the exterior as male pore (see Image 2 and Image 4). Lone et al. (2022) reported high intra specific divergence for M. travancorensis. Female pores also clearly demarcated on 13 (Image 5A), however, not much ornamented and associated setae at ab (Image 5B). Because of its less physical diversity across species and relatively conserved position, the female pore is often regarded as having minimal taxonomic relevance. This work provides important taxonomic and systematic insights into M. travancorensis’s distinctive reproductive morphology by highlighting the genital markings and related structures in and around the clitellum.

 

 

CONCLUSION

 

Highly specialized and decorated genital characteristics that are important for species identification and reproductive function are revealed by the ultrastructural investigation of M. travancorensis. The male genital region is complex, as indicated by the existence of paired prostatic ducts, a seminal groove, and eye-shaped genital marks. Small structural adaptations such as retracted penial setae, lateral prostatic pores, seminal grooves, and glandular papillae can be found with SEM imaging. These discoveries provide useful taxonomic markers within the genus Megascolex and advance our knowledge of earthworm reproductive characteristics. Moreover, the distinct genital specializations observed may represent adaptations for efficient sperm transfer, mate recognition, and reproductive isolation, thereby offering insights into phylogenetic relationships.

 

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REFERENCES

 

Ambreen, F., S. Lubna, A. Shamim & M. Yasmin (2004). Study of Genital Variations among the Earthworms of Genus Pheretima Inhabiting the Soil of Islamabad. Pakistan Journal of Biological Science 7: 123–125. https://doi.org/10.3923/pjbs.2004.123.125

Aiyer, K.S.P. (1929). Oligochaetes of Travancore. D.Sc. Thesis. University of Madras, India.

Beddard, F.E. (1895). A Monograph of the Order Oligochaeta. Oxford: Clarendon Press, xxi + 769 pp.

Blakemore, R.J. (2006). Cosmopolitan Earthworms an Eco-taxonomic guide to the peregrine species of the World. Verm Ecology (ed.), 2nd edition, Yokohama, Japan, 600 pp.

Caramelo, C. & E. Martínez-Ansemil (2012). Morphological investigations of microdrile oligochaetes (Annelida, Clitellata) using scanning electron microscopy. Turkish Journal of Zoology 36: 1–14. https://doi.org/10.3096/zoo-1002-25.

Erséus, C. (1981). Taxonomic studies of Phallodrilinae (Oligochaeta, Tubificidae) from the Great Barrier Reef and the Comoro Islands, with descriptions of ten new species and one new genus. Zoologica Scripta 10(1): 15–31. https://doi.org/10.1111/j.1463-6409.1981.tb00481.x

Cosín, D.J.D., M. Novo & R. Fernández (2011). Reproduction of Earthworms: Sexual Selection and Parthenogenesis, pp. 69–86. In: Karaca, A. (ed.). Biology of Earthworms, Soil Biology 24. Springer, Berlin, Heidelber. https://doi.org/10.1007/978-3-642-14636-7_5

Gates, G.E. (1972). Burmese Earthworms, an introduction to the systematics and biology of Megadrile oligochaetes with special reference to South-East Asia. Transactions of the American Philosophical Society (n.s.), 62(7): 1–326.

Gates, G.E. & J. W. Reynolds (2017). Preliminary Key to North American Megadriles (Annelida, Oligochaeta), based on external characters, insofar as possible. Megadrilogica 22(10): 201–219.

Kamal S., N. Chachain & M. Sabbah (2023). The Morphological identification of earthworms in Baghdad Governate. Envirionmental Science, Biology Sumer 2(8): 1. https://doi.org/10.21931/RB/CSS/2023.08.02.35

Khan, M.U., S. Andleeb & M.F. Khan (2022). Biodiversity and ecological interactions of earthworm species from Poonch division Pakistan. Tropical Ecology 63: 122–133. https://doi.org/10.1007/s42965-021-00196-9.

Jamieson, B.G.M. (2001). Native Earthworms of Australia (Megascolecidae, Megascolecinae). Supplement, August 2001. Science Publishers, Inc., Enfield, New Hampshire, USA (Compact disc).

Julka, J.M. (1988). The Fauna of India and Adjacent Countries, Megadrile Oligochaeta (Earthworms), Haplotaxida: Lumbricina: Megascolecoidea: Octochaetidae. Zoological Survey of India, Calcutta, 400 pp.

Michaelsen, W. (1900). Oligochaeta. Lief. 10. In: Das Tierreich. Berlin: Friedländer, xxix + 575 pp.  https://doi.org/10.5962/bhl.title.11605

Michaelsen, W. (1907). Oligochaeta. (Fauna Sudwest-Australiens, hrsg. V.W. Michaelsen und R. Hartmeyer, Bd. 1, Lfg.2) Jena: G. Fischer 1907: 117–232.

Michaelsen, W. (1908). The Oligochaeta of India, Nepal, Ceylon, Burma and the Andaman Islands. Memoirs of the Indian Museum, Calcutta 1(3): 103–253. https://www.biodiversitylibrary.org/part/92322.

Michaelsen, W. (1910). Die oligochätenfauna der Vorderindisch-Ceylonischen Region. Abhandlingen aus dem Gebiete der naturwissenschaften hrsg. vom naturwissenschaftlichen Verein in Hamburg 19(5): 1–108.

Michaelsen, W. (1913). Oligochäten von Travancore und Borneo. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Hamburg 30(2): 73–92.

Michaelsen, W. (1916). Results of Dr. E. Mjöberg’s Swedish Sacientific Expedition to Australia 1910-1913. XIII. Oligochäten. Kungliga Svenska Vetenskapsakademiens Handlingar, Stockholm 52(13): 1–74.

Olive, P.J.W. & R.B. Clark (1978). Physiology of reproduction. pp. 271–368. In: Mill, P.J. (ed.). Physiology of Annelids. Academic Press, London, v + 710 pp.

Narayanan, S.P., R. Paliwal, A. Thomas & J.M. Julka (2025). Current Status of Earthworm Fauna (Clitellata: Megadrili) of Parambikulam Tiger Reserve, within the western Ghats of Kerala, India. Journal of the Bombay Natural History Society (JBNHS) 122(1) https://doi.org/10.17087/jbnhs/2025/v122/165060  

Premachandran, P.N. (ed.) (2007). Bench Mark Soils of Kerala. Soil Survey Organisation Agriculture Department, Government of Kerala, Department of Agriculture, Kerala, South India, 588 pp.

Ramasamy, P.K. (2009). Biodiversity of Earthworms in the Eastern Ghats of (Sathyamangalam Division) Tamilnadu, India and Influence of Vermicompost on Growth, Yield and Nutritional Status of Some Selected Plants. Ph.D. Thesis, Bharathiar University. Tamil Nadu, India.

Rathour, S. & J.M. Julka (2024). Monthly variation of Diversity, Density and Biomass of Earthworm in Mid Hills of North Western Himalayas under Three Different Land Use Systems, District Bilaspur, Himachal Pradesh African Journal of Biological Science 6(5): 6315–6337.

Reynolds, J.W. (1974). Are Oligochaetes really hermaphroditic amphimictic organisms? The Biologist 56(2): 90–99.

Reynolds, J.W. (2022). The earthworms (Lumbricidae, Megascolecidae and Sparganophilidae) in Canada.  Ottawa: Canadian Food Inspection Agency, i + 179 pp.

Reynolds, J.W. & M.J. Wetzel (2025). Nomenclatura Oligochaetologica – A catalogue of names, descriptions and type specimens of the Oligochaeta. Editio Secunda. URL: https://nomenclatura-oligochaetologica.inhs.illinois.edu  Accessed on 23.ix.2025.

Rosa, D. (1891). Die exotichen Terricolen des K.K. naturhistorischen Hofmuseum. Annale naturhistorischen museum, Wien 6 (3&4): 379–406.

Stephenson, J. (1923). Oligochaeta. Fauna of British India including Ceylon and Burma. London: Taylor and Francis, xxiv + 518 pp.

Templeton, R. (1844). Description of Megascolex caeruleus. Proceedings of the Zoological Society of London 12 (1844): 89–91.

Yáñez, E., S. Cuadrado & E. Martínez-Ansemil (2006). External sense organs in freshwater oligochaetes (Annelida, Clitellata) revealed by scanning electron microscopy Journal of Morphology 267: 198–207. https://doi.org/10.1002/ jmor.10397