Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 May 2026 | 18(5): 28838–28844
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.8978.18.5.28838-28844
#8978 | Received 18 February 2024 | Final received 16 January 2026|
Finally accepted 18 March 2026
Eastern range record of the
semiaquatic freshwater earthworm Glyphidrilus
gangeticus Gates, 1958 (Clitellata:
Crassiclitellata: Almidae)
from West Bengal, India, with a brief key to the Indian species of the genus
M. Nurul Hasan 1 , John Warren Reynolds 2 , Hasko F. Nesemann
3, Shyamasree Ghosh 4
1,⁵ Zoological Survey of India
(Ministry of Environment, Forest and Climate Change, Government of India), FPS
Building, Indian Museum Campus, 27 JL Nehru Road, Kolkata, West Bengal 700016,
India.
² Oligochaetology Laboratory, Kitchener, Ontario, N2A 4E1
Canada and New Brunswick Museum, Saint John, New Brunswick, E2K 1E5, Canada.
³ Im Obergarten 9, D-65719 Hofheim am
Taunus, Hesse, Germany.
⁴ School of Biological
Sciences, National Institute of Science Education and Research (NISER),
Bhubaneswar, Odisha, India.
4 Homi Bhabha National Institute,
Training School Complex , Anushakti
Nagar, Mumbai 400094, India.
1 nhasan.mld@gmail.com
(corresponding author), 2 john.w.reynolds1941@gmail.com, 3 hnesemann2000@yahoo.co.in,
4 shyamasree_b@yahoo.com, 5 mandalsucker@gmail.com
Abstract: The semi-aquatic earthworm
genus Glyphidrilus is poorly
documented in West Bengal, India, with only one species previously recorded. To
fill this distribution gap, we surveyed riverbanks of the Mahananda River (West
Bengal) and Ganga River (Bihar). Earthworms were collected by hand sorting
from wet soil and mud during low-water conditions in 2021 and 2024, preserved
in 4% formalin, and identified using morphological characters under a
stereomicroscope. We report the first record of Glyphidrilus gangeticus Gates,
1958 from West Bengal, extending its known range eastward from Uttar Pradesh
and Bihar. Additionally, we provide a revised identification key to all seven
Indian species of Glyphidrilus (annandalei, elegans,
fluviatilis, gangeticus,
papillatus, spelaeotes,
and tuberosus) based on literature and newly
examined material. This discovery increases the number of Glyphidrilus species known from West Bengal to
two and highlights the need for further surveys of semi-aquatic habitats in the
Gangetic plains. The improved key will aid future biodiversity assessments and
conservation planning.
Keywords: Almidae,
first record, freshwater biodiversity, Ganga River, identification key,
Mahananda River, semiaquatic earthworms.
Editor: Bruce Snyder, Georgia College & State
University, Milledgeville, Georgia. Date
of publication: 26 May 2026 (online & print)
Citation: Hasan,
M.N., J.W. Reynolds, H.F. Nesemann, S. Ghosh &
C.K. Mandal (2026). Eastern range record of the semiaquatic freshwater earthworm Glyphidrilus gangeticus
Gates, 1958 (Clitellata: Crassiclitellata:
Almidae) from West Bengal, India, with a brief key to
the Indian species of the genus. Journal of
Threatened Taxa 18(5):
28838–28844. https://doi.org/10.11609/jott.8978.18.5.28838-28844
Copyright: © Hasan 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: None.
Competing interests: The authors declare no competing interests.
Author details: M. Nurul Hasan, Ph.D. research scholar, is working on the annelid fauna of India with special emphasis on taxonomy and biodiversity studies. He is affiliated with the Zoological Survey of India (Ministry of Environment, Forest and Climate Change, Government of India), West Bengal. John Warren Reynolds, Ph.D., LL.B., is director of the Oligochaetology Laboratory, Kitchener, Canada, and research associate at the University of New Brunswick, Saint John, Canada. His specialization is the taxonomy, systematics, and biogeography of the Oligochaeta. Hasko F. Nesemann, Ph.D., is an Independent Researcher affiliated with Im Obergarten 9, Germany. His research interests include freshwater biodiversity, aquatic ecology, and annelid taxonomy. Shyamasree Ghosh, Ph.D., is working in the capacity of Scientific Officer at the School of Biological Sciences (SBS), National Institute of Science Education and Research, Bhubaneswar, Odisha, India, under the Homi Bhabha National Institute, Mumbai. Her research activities involve molecular biology and biodiversity sciences. Chandra Kanta Mandal, Ph.D., scientist, is an expert on the annelid fauna of India and is affiliated with the Zoological Survey of India (Ministry of Environment, Forest and Climate Change, Government of India), West Bengal.
Author
contribution: MNH, JWR and CKM: conceptualization, investigation and writing – original draft. MNH, JWR, CKM and SG: data curation, monitoring and validation. MNH, JWR, CKM and SG: visualization. MNH, JWR, CKM, SG and HN: writing, reviewing and editing. All authors contributed to the article and approved the submitted version.
Acknowledgments: The authors acknowledge the director, Zoological Survey of India (ZSI, India), and the director, NISER Bhubaneswar, for the study. We also thank the reviewers for their comments, which significantly improved the manuscript.
Introduction
There are approximately 5,739
species and subspecies of megadrile earthworms worldwide (Marchán
et al. 2022; Mısırlıoǧlu et al. 2023; Reynolds &
Wetzel 2025). At present, India has 457 species under 10 families – Almidae, Acanthodrilidae, Benhamiidae, Eudrilidae, Lumbricidae, Moniligastridae, Megascolecidae, Ocnerodrilidae, Octochaetidae, and Rhinodrilidae
(Balakrishnan et al. 2024). India hosts 12.5% of the world’s known earthworm
species (Julka et al. 2009), with 71% of genera and
85% of species endemic (Julka & Paliwal 2005). Glyphidrilus (40
species-level taxa) is found in China, Cambodia, India, Indonesia, Laos,
Myanmar, Sri Lanka, Singapore, and Thailand, but only one species, G. stuhlmanni Michaelsen,
1897, has been recorded from Africa (Chanabun et al.
2013). Glyphidrilus is mainly
represented in the Gangetic plains and associated freshwater habitats.
Classical works by Stephenson (1923) and Julka
(1988) documented the presence of several species, including G. gangeticus, G. tuberosus, G.
fluviatilis, G. elegans, G.
spelaeotes, G. papillatus,
and G. annandalei. These records
highlight the genus as an important faunal component of Indian freshwater
ecosystems. Recent surveys and taxonomic contributions (Narayanan et al. 2023)
continue to confirm the occurrence of these species, while also emphasizing the
need for detailed ecological and molecular studies to resolve species
boundaries.
Despite these records, the
earthworm fauna of West Bengal remains poorly explored for semi-aquatic Glyphidrilus species. Only G. tuberosus has been reported from the state (Ahmed
et al. 2022), leaving a significant distributional gap for other species,
including G. gangeticus, which is known
from adjacent states (Uttar Pradesh and Bihar).
We hypothesize that Glyphidrilus gangeticus occurs
in West Bengal due to the hydrological continuity of the Ganga River system,
which may facilitate eastward dispersal along the Gangetic floodplains.
The present study was undertaken
to: (1) survey selected riverbanks in West Bengal and Bihar for the presence
of Glyphidrilus species; (2) confirm
the identity of any newly collected material using morphological characters;
(3) report the first record of G. gangeticus from
West Bengal, if found; and (4) compile an improved identification key for all
seven Indian Glyphidrilus species to
aid future research.
Glyphidrilus species play important
roles in sediment aeration and nutrient cycling and can serve as bioindicators
of freshwater ecosystem health (Chanabun et al.
2013). Detailed ecological functions have been reviewed elsewhere and are
not repeated here to avoid redundancy. Many regions of India remain
unexplored for earthworms, and targeted surveys are needed to fill knowledge
gaps. The present study addresses this need by providing a new state record and
a practical identification tool.
Materials
and Methods
The study areas were the
semiaquatic riverbanks of the lower Mahananda River in West Bengal and the
Ganga River in Bihar (Figure 1).
The earthworms were collected in
low-water conditions during the years 2021 and 2024 from the wet soil and mud
by digging and sorting by hand from fallen tree trunks and leaf litter. The
collected samples of earthworms were washed with water and preserved in 4%
formalin overnight and then transferred to 70% alcohol for identification.
Identification was done using a Leica EZ4HD stereomicroscope. The morphometric
characters of the species were carefully recorded after preservation.
Measurements were taken manually using a slide caliper, which allowed accurate
determination of body length, width, and segmental details. This method ensured
precision in documenting the diagnostic features essential for taxonomic study.
The collected specimens were identified using Blakemore (2012), Gates (1972), Julka (1988), and Stephenson (1923). The specimens were
deposited with the Zoological Survey of India, headquarters, Kolkata (ZSI), and
additional specimens at the Gangetic Plains Regional Centre, Patna.
Results
Genus Glyphidrilus
Horst, 1889
Glyphidrilus Horst, 1889: 86. Type species: Glyphidrilus weberi
Horst, 1889.
Bilimba Rosa, 1890: 386.
Annadrilus Horst, 1893: 44.
Glyphidrilus + Bilimba
+ Annadrilus - Beddard,
1895: 679, 686, 680.
Glyphidrilus Michaelsen,
1900: 459.
Glyphidrilus Michaelsen,
1909: 244.
Glyphidrilus Michaelsen,
1910: 99.
Glyphidrilus Michaelsen,
1918: 343.
Glyphidrilus Gates, 1972: 234.
Currently, 40 species of the Glyphidrilus genus have been reported from Asia
(from the Indonesian islands to the Malay Peninsula and Myanmar, extending to
western India, to northern China and Africa (Gates 1951; Naidu 2005; Chanabun et al. 2013). Thus, a key for seven species of Glyphidrilus found in India is presented. These taxa
are briefly characterized by their external features together with a short
synonymy, based on the cited references.
1. Glyphidrilus
gangeticus Gates, 1958
Glyphidrilus sp.? Michaelsen,
1909: 244.
Glyphidrilus papillatus Stephenson, 1920: 258–260.
Glyphidrilus papillatus (part) - Stephenson, 1923: 493.
Glyphidrilus sp. - Gates, 1947: 121.
Glyphidrilus papillatus Gates, 1948: 175–176.
Glyphidrilus papillatus Gates, 1951: 17.
Glyphidrilus gangeticus Gates, 1958: 58 Type missing
(Reynolds & Wetzel, 2025).
Glyphidrilus gangeticus Narayanan et al., 2023: 114.
Type locality: Saharanpur, Uttar Pradesh State,
India.
Material examined: (from Gates 1959; Stephenson
1920). India; West Bengal, Mahananda River near Telaigachi
Village, Chanchal, Malda, 25.302° N, 88.111° E, 6 m;
26.xii.2021; M. Nurul Hasan leg.; 9 clitellate adults (0-0-9) ex. Reg. no.
ZSI-An6247/1. Bihar State, Ganga River downstream Patna, Ghaighat,
25.606° N, 85.221° E, 64 m; 12.iv.2024, Gopal Sharma with Hasko
Nesemann, 2 clitellate adults (0-0-2) ex. Reg. no.
NDRC/IV/2024.
Brief description: Length 85–140 mm, diameter 3–5
mm. Number of segments 202–340. The prostomium is zygolobous.
Clitellum, saddle, xiii, xvi, xvii–xxxiv. Wings xviii–xxiii. Spermathecal pores
are not visible in 12/13–16/17 segments. One female pore on 14 near setal line B, about midway between 13/14 and the setal equator. Dorsal pores absent. The G. gangeticus morphological characters of the gizzard, heart,
seminal vesicles (4 pairs), spermathecae (4 pairs), nephridia and dorsal blood
vessels are provided in Image 1. Setae closely paired. Prostates absent.
Typhlosole begins in xviii.
Distribution: India (Assam, Delhi,
Uttarakhand, Uttar Pradesh, Bihar (Michaelsen 1909;
Stephenson 1920; Gates 1951; Naidu 2005; Chanabun et
al. 2013), and West Bengal); Thailand (Nesemann et
al. 2004, 2007).
2. Glyphidrilus
tuberosus Stephenson, 1916
Glyphidrilus tuberosus, Stephenson, 1916: 349. Type:
Zoological Survey of India, Calcutta, cat. no. 6517 (Reynolds & Wetzel
2025).
Glyphidrilus papillatus Gates, 1958: 59.
Glyphidrilus tuberosus Narayanan et al., 2023: 114.
Type locality. Kendrapara
Canal near Cuttack, Odisha State, India.
Material examined: None.
Brief description: (from Gates 1958; Stephenson
1916). Length 60 mm, maximum diameter 3 mm. Number of segments 215–221. Size of
the segments is smaller after the clitellar region. Colour
is brown. Dorsal surface is concave in arrears of the clitellum xiv, xv,
xvi–xxviii, xxix, xxx, ventral surface flat and concave occasionally. Wings
xx–xxiv. The section of the body looks four-sided. Prostomium pro- or zygolobous. Setae are widely paired. Gizzard in vii. No
calciferous glands.
Distribution: India (Bihar, Jharkhand,
Karnataka, Madhya Pradesh, Odisha, Tamil Nadu, Tripura, Uttarakhand, Uttar
Pradesh and West Bengal (Stephenson 1923, Julka &
Senapati 1987 and Mandal & Kumar 2018)); Bangladesh (Das & Reynolds
2003).
3. Glyphidrilus
annandalei Michaelsen,
1910
Glyphidrilus annandalei Michaelsen,
1910: 101. Type: Zoologisches Museum Universität
Hamburg cat. no. 3600! (Reynolds & Wetzel, 2025)
Glyphidrilus annandalei Cognetti,
1900: 502.
Glyphidrilus achencoili (laps.) - Michaelsen,
1913: 92.
Glyphidrilus annandalei - Stephenson, 1916: 349.
Glyphidrilus annandalei- Michaelsen,
1918: 344.
Glyphidrilus annandalei - Stephenson, 1921: 767.
Glyphidrilus fluciatilis + G. elegans
+ G. rarus + G. saffronensis
- Rao, 1922: 53, 66.
Glyphidrilus annandalei - Stephenson, 1922: 387.
Glyphidrilus annandalei - Narayanan et al., 2023: 112.
Type locality: Quilon (Kollam), Kerala State,
India.
Material examined: None.
Brief description: (from Michaelsen,
1910; Stephenson, 1916, 1921). Length 90–265 mm, diameter 2.5–11 mm. Number of
segments 125–322. Colour dark grey, without any
pigmentation; a slight pink colour is found on the
clitellum. Four-sided section found in its subsequent half of the body. The
dorsal side broader than the ventral side.
Distribution: Endemic to India (Karnataka,
Kerala (Rao 1922; Chanabun et al. 2013; Narayanan et al. 2023), Madhya
Pradesh, Tamil Nadu, Uttarakhand, Uttar Pradesh (Stephenson 1923)).
4. Glyphidrilus
elegans Rao, 1922
Glyphidrilus elegans Rao, 1922; 62. Type locality:
Mysore, India. Type: British Museum (Natural History) 1922:4:20:9! (Reynolds
& Wetzel, 2025).
Glyphidrilus annandalei - Stephenson, 1923: 491.
Glyphidrilus annandalei Brinkhurst
& Jamieson, 1971: 755.
Glyphidrilus elegans Chanabun
et al., 2013: 27.
Glyphidrilus elegans - Narayanan et al., 2023: 113.
Type locality: Sandy islets in the Cauvery
River in Dubari Forest at Fraserpett
(=Kushalnagar), Karnataka State, India.
Material examined: None.
Brief description: (from Rao 1922; Stephenson 1923;
Brinkhurst & Jamieson 1971). Length 139 mm. Body
quadrangular in transverse section behind the clitellum, cylindrical in the
anterior region. Total segments 248. Body tone light brown. Dorsal surface
noticeably wider than the ventral surface at the posterior end. In the ventro-lateral region of the clitellum, setae are seen in
25–31.
Distribution: India (Karnataka, Tamil Nadu (Kathireswari et al. 2008), Rajasthan, Punjab, Haryana, and
Gujarat (Dhiman & Battish
2006)); Bangladesh; Myanmar (Reynolds et al. 1995 as G. e. ssp. paski Stephenson, 1923); Thailand (Chanabun et al. 2013).
5. Glyphidrilus
fluviatilis Rao, 1922
Glyphidrilus fluviatilis Rao, 1922: 53. Type: British
Museum (Natural History) 1922:4:20:5! (Reynolds & Wetzel, 2025).
Glyphidrilus annandalei
Stephenson,
1923: 481.
Glyphidrilus annandalei Brinkhurst
& Jamieson, 1971: 755.
Glyphidrilus fluviatilis
Chanabun et al., 2013: 27.
Glyphidrilus fluviatilis Narayanan et al., 2023: 113.
Type locality: Sandy banks of the Harangi River in Madapur,
Telangana State, India.
Material examined: None.
Brief description: (from Rao 1922). Length 272 mm.
Body quadrangular behind the clitellum, anterior half cylindrical. Number of
segments 225.
Distribution: India (Karnataka); Thailand (Chanabun et al. 2013).
6. Glyphidrilus
papillatus (Rosa, 1890)
Bilimba papillata Rosa, 1890: 386.
Bilimba papillatus - Beddard,
1895: 687.
Glyphidrilus papillatus Michaelsen,
1896: 196.
Glyphidrilus papillatus Michaelsen,
1900: 459.
Glyphidrilus papillatus
Michaelsen, 1918: 344.
Glyphidrilus papillatus Stephenson, 1920: 58.
Type locality: Cobapo
(Carin Mountains), Burma (Myanmar).
Material examined: None.
Brief description: (from Stephenson 1920). Body
length 74–120 mm, maximum diameter 3–6 mm. Total number of segments 130–330. No
pigmentation dorsally, colour greyish. Secondary
annulation found after five segments of the anterior part. Posterior half
flattened dorsally. Prostomium pro- or zygolobous.
Dorsal pores absent. Clitellum, saddle xiii, xiv–xl. Wings begin in xvii–xix
and end in xxiii–xxvii. Genital markings, unpaired and median in xi–xxi, paired
and lateral to A in xii–xxx. Gizzard in viii. Intestinal origin in xv.
Typhlosole, lamelliform beginning in xviii.
Distribution: India (Uttar Pradesh (Stephenson
1923)); Myanmar (Stephenson 1923).
7. Glyphidrilus
spelaeotes Stephenson, 1924
Glyphidrilus spelaeotes Stephenson, 1924: 133. Type:
Zoological Survey of India, Calcutta, cat. no. 1155–1156 (Reynolds &
Wetzel, 2025).
Glyphidrilus papillatus (part.) Gates, 1958: 54.
Glyphidrilus papillatus Brinkhurst
& Jamieson, 1971: 764.
Glyphidrilus spelaeotes Narayanan et al., 2023: 114.
Type locality: Siju
Cave, Meghalaya State, India.
Material examined: None.
Brief description: (from Stephenson 1924; Brinkhurst & Jamieson 1971). Length 175 mm, diameter
2–3 mm, segment number 310. Pale or light brownish-grey colour,
four-sided behind the wings. The wings are attached between the line of the
dorsal and that of the ventral setal couples, and extend from xviii or xix to xxiv or xxv; they
are bent downwards and inwards, and on segment xviii they are, if present, less
prominent than in the rest of their extent.
Distribution: India: Assam, Meghalaya
(Stephenson 1924).
Discussion
Glyphidrilus are restricted to subtropical
Africa and southern Asia (Magalhães et al. 2021).
These semi-aquatic earthworms usually inhabit river banks, wetlands, and muddy
substrates, where they play an important role in soil aeration and nutrient
cycling. Their restricted distribution makes them ecologically significant
indicators of freshwater-associated habitats. Many regions of India are still
unexplored in terms of earthworm fauna. Extensive surveys have been conducted
in some biodiversity-rich zones, but several river systems, wetlands, and
floodplains remain poorly studied. This gap in knowledge highlights the
possibility of discovering new records or even undescribed species, especially
in areas with unique hydrological conditions. From West Bengal, only Glyphidrilus tuberosus
was known (Ahmed et al. 2022). This species was previously reported from the
Ganga-Brahmaputra Basin and is characterized by its distinct gizzard position
and genital markings. Glyphidrilus gangeticus is the first record of this species from this
state. Earlier, this species was reported from Uttar Pradesh and Bihar along
the Ganga River system, and its occurrence in West Bengal extends the known
distribution range eastward. The finding demonstrates the connectivity of
aquatic habitats across the Ganga floodplains and lack of systematic studies.
This list summarizes all known species from India with details of their
diagnostic characters, type localities, and ecological preferences. Glyphidrilus gangeticus
is the second species from this state. Thus, the state now contributes two
members of this semi-aquatic genus, enriching the faunal diversity of the
region. There are seven species of the genus Glyphidrilus
known from India. These include G. tuberosus, G.
gangeticus, and five others described from
various river basins across Uttar Pradesh, Bihar, Assam, and Meghalaya. Their
distributions are often linked to major river systems such as the Ganga,
Brahmaputra, and Barak. This discovery is significant as it not only adds to
the state checklist but also provides baseline information for future
biodiversity assessments and conservation planning of semi-aquatic habitats.
The study emphasizes the need for a taxonomy combining morphology with
molecular data. The local faunistic studies largely contribute to our knowledge
of poorly known semi-aquatic species. Such studies are crucial in recognizing
habitat-specific taxa, detecting biogeographical patterns, and informing
conservation priorities in freshwater ecosystems.
Key to the Indian species of the
genus Glyphidrilus Horst, 1889
1. Clitellum begins on segment 13
..............................................................................................................
2
1’. Clitellum begins on segment 14
..............................................................................................................
5
2. Clitellum on 13–34; wings on segments 17–25
.................................................................... G. gangeticus
2’. Wings begin on segment 25
.....................................................................................................................
3
3. Wings extend beyond segment 32; wings on segments 25–36;
clitellum beginning 13–18, ending 35–41;
spermathecal
pores in 13/14–16/17 ....................................................................................
G. annandalei
3’. Wings end on or before segment 32
.........................................................................................................
4
4. Wings on segments 25–31; clitellum on 13–35; spermathecal pores
in 13/14–17/18 ............... G. elegans
4’. Wings on segments 25–32; clitellum on 13–33 (occasionally extends
to 36 or 38); spermathecal pores in
intersegmental
furrows 7/8 and 8/9
.........................................................................................
G. fluviatilis
5. Wings on segments 20–24 to 28; clitellum on 14–30; spermathecal
pores in 14–15 ............ G. tuberosus
5’. Wings begin before segment 20
..............................................................................................................
6
6. Wings on segments 18–25; clitellum on 14–30; spermathecal pores
in 13/14–15/16 ......... G. spelaeotes
6’. Wings on segments 18–26; clitellum on 14–40; spermathecal
pores in 14–17 ...................... G. papillatus
For
figure & image - - click here for full PDF
References
Ahmed, S., P.
Raj & D. Gupta (2022). Annelida: Clitellata: Oligochaeta
and Hirudinea, pp. 85–90. In: Chandra, K., D.
Banerjee, C. Raghunathan, D. Gupta, P. Raj & D. Sharma (eds.). Faunal
Diversity of Biogeographic Zones of India: Gangetic Plains. Zoological
Survey of India, Kolkata.
Beddard, F.E. (1895). A Monograph of the Order Oligochaeta. Clarendon Press, Oxford, xii + 769 pp.
Brinkhurst, R.O. & B.G.H. Jamieson
(1971). Aquatic Oligochaeta of the World. University of Toronto Press,
Toronto, xi + 890 pp.
Blakemore,
R.J. (2012). Cosmopolitan
Earthworms, 5th Edition. Verm Ecology, Yokohama. CD-ROM Monograph, 990 pp.
Balakrishnan,
S., T. Mondal, C.K. Mandal, S. Ahmed, J.S.W. Kumar, G. Sivaleela
& C. Raghunathan (2024). Fauna of India Checklist: Annelida: Polychaeta:
Sipuncula: Clitella: Hirudinida: Crassiclitellata:
Moniligastrida. Online Version 1.0. Zoological
Survey of India, Kolkata. https://zsi.gov.in/uploads/documents/checklist/english/026_Annelida.pdf.
Accessed on 20.iv.2026.
Cognetti de Martiis,
L. (1900). Contributo alla conoscenza degli Oligocheti neotropicali. Bollettino dei musei di zoologia ed anatomia comparata
della R. Università di
Torino 15(369): 1–17.
Chanabun, R., C. Sutcharit,
P. Tongkerd & S. Panha
(2013). The
semi-aquatic freshwater earthworms of the genus Glyphidrilus
Horst, 1889 from Thailand (Oligochaeta: Almidae) with redescriptions of
several species. ZooKeys 256: 1–76.
Das, B.C.
& J.W. Reynolds (2003). A checklist and distribution of the freshwater and terrestrial Annelida
(Oligochaeta, Hirudinea and
Polychaeta) of Bangladesh. Megadrilogica
9(10): 61–70.
Dhiman, N. & S.K. Battish (2006). Earthworms from northern Indian
states with Ocnerodrilus occidentalis
Eisen, 1878 as a new report from Punjab. Zoos’ Print Journal 21(1):
2135–2137. https://doi.org/10.11609/JoTT.ZPJ.1203.2135-7
Gates, G.E.
(1947). Earthworms
of the Allahabad sector of the Gangetic Plain. Proceedings of the National
Academy of Science India, Allahabad 17: 117–128.
Gates, G.E. (1948). On some earthworms from the Buitenzorg
Museum. III. Results of the Third Archbold Expedition 1938–1939. Treubia, Buitenzorg
19: 139–166.
Gates, G.E. (1951). On the earthworms of Saharanpur, Dehra Dun, and
some Himalayan hill stations. Proceedings of the National Academy of Science
India, Allahabad (B) 21: 16–22.
Gates, G.E. (1958). On Indian and Burmese earthworms of the genus Glyphidrilus. Records of the Indian Museum 53(1–2):
53–66.
Gates, G.E. (1972). Burmese earthworms. An introduction to the
systematics and biology of megadrile oligochaetes with special reference to
Southeast Asia. Transactions of the American Philosophical Society, n.s. 62(7): 1–326. https://doi.org/10.2307/1006214
Hasan, M.N., S. Ahmed, K. Deuti & N. Marimuthu (2023). Earthworm
(Annelida: Clitellata) fauna of Chhattisgarh, India. Journal
of Threatened Taxa 15(4): 23091–23100. https://doi.org/10.11609/jott.8135.15.4.23091-23100
Horst, R. (1889). Over eene nieuwe soort der Lumbricinen door Prof. Max Weber uit
Nederl. Indie medegebracht.
Tijdschrift der Nederlandsche
Dierkundige Vereeniging 2: 1–77.
Horst, R. (1893). Earthworms from Malay Archipelago. In:
Weber, M. (ed.). Zoologische Ergebnisse
einer Reise in Niederländisch Ost-Indien 3:
28–83.
Julka, J.M.
(1988). The Fauna of India and Adjacent Countries: Megadrile Oligochaeta (Earthworms). Haplotaxida:
Lumbricina: Megascolecoidea:
Octochaetidae. Zoological Survey of India, Calcutta,
xiv + 400 pp.
Julka, J.M. &
R. Paliwal (2005).
Distribution of earthworms in different agro-climatic
regions of India, pp. 3–13. In: Ramakrishnan, P.S., K.G. Saxena, M.J. Swift,
K.S. Rao & R.K. Maikhuri (eds.). Soil
Biodiversity, Ecological Processes and Landscape. Oxford and ABH
Publications, New Delhi, 462 pp.
Julka, J.M. &
B.P. Senapati (1987). Earthworms (Oligochaeta:
Annelida) of Orissa, India. Zoological Survey of India, Miscellaneous
Publication, Occasional Paper, No. 92, 49 pp.
Julka, J.M., R. Paliwal & P. Kathireswari
(2009). Biodiversity of Indian earthworms – an overview, pp. 36–56. In: Edwards,
C.A., R. Jayaraaj & I.A. Jayaraaj
(eds.). Proceedings of the Indo-US Workshop on Vermitechnology
in Human Welfare. Rohini Achagam, Coimbatore, India,
108 pp.
Mandal, C.K. & H. Kumar (2018). Annelida: Oligochaeta, pp. 139–144. In: Chandra, K., D. Gupta, K.C.
Gopi, B. Tripathy & V. Kumar (eds.). Faunal
Diversity of Indian Himalaya. Zoological Survey of India, Kolkata.
Marchán, D.F., S.W.
James, A.R. Lemmon, E.M. Lemmon, M. Novo, J. Domínguez, D.J.D. Cosín & D. Trigo (2022). A strong
backbone for an invertebrate group: anchored phylogenomics
improves the resolution of genus-level relationships within the Lumbricidae (Annelida, Crassiclitellata).
Organisms Diversity & Evolution 22: 915–924.
Michaelsen, W. (1896). Oligochaeten. Ergebnisse der zoologischen Forschungsreise in
den Molukken und in Borneo (Kükenthal).
Abhandlungen herausgegeben
von der Senckenbergischen Naturforschenden
Gesellschaft, Frankfurt a. Main 23: 193–243.
Michaelsen, W. (1900). Das Tierreich. 10. Lieferung: Oligochaeta. R. Friedländer und
Sohn, Berlin, xxix + 575 pp.
Michaelsen, W. (1909). The Oligochaeta of India, Nepal, Ceylon, Burma and the Andaman
Islands. Memoirs of the Indian Museum, Calcutta 1(3): 103–253.
Michaelsen, W. (1910). Die Oligochätenfauna der vorderindisch-ceylonischen
Region. Abdruck aus
den Verhandlungen des Naturwissenschaftlichen
Vereins 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. (1918).
Die Lumbriciden, mit
besonderer Berücksichtigung
der bisher als Familie Glossoscolecidae zusammengefassten Unterfamilien. Zoologische Jahrbücher,
Abteilung für Systematik, Jena 41: 1–398.
Mısırlıoğlu, M., J.W.
Reynolds, M. Stojanović, T. Trakić,
J. Sekulić, S.W. James, T. Decaëns,
E. Lapied, H. Phillips, E. Cameron & G.G. Brown
(2023). Earthworms (Clitellata, Megadrili)
of the world: an updated checklist of valid species and families, with notes on
their distribution. Zootaxa 5255(1): 417–438. https://doi.org/10.11646/zootaxa.5255.1.33
Naidu, K.V. (2005). The Fauna of India and the Adjacent
Countries – Aquatic Oligochaeta. Zoological
Survey of India, Kolkata, 294 pp.
Narayanan, S.P., R. Paliwal, S. Kumari, S. Ahmed, A.P. Thomas & J.M. Julka (2020). Annelida: Oligochaeta,
pp. 87–102. In: Faunal Diversity of Biogeographic Zones of India: Western
Ghats. Zoological Survey of India, Kolkata, 744 pp.
Narayanan, S.P., R. Paliwal, V.T. Kurien, A.P. Thomas & J.M. Julka
(2023). Earthworms of India (Clitellata: Moniligasteridae, Crassiclitellata):
Distribution and Status. Mahatma Gandhi University, Kottayam, ix + 378 pp.
Narayanan, S.P., S. Sathrumithra, R. Anuja, G. Christopher, A.P. Thomas & J.M. Julka (2021). Three new species and four new
species records of earthworms of the genus Moniligaster
Perrier, 1872 (Clitellata: Moniligastridae)
from Kerala region of the Western Ghats biodiversity hotspot, India. Zootaxa 4949(2): 381–397. https://doi.org/10.11646/zootaxa.4949.2.11
Narayanan, S.P., V.T. Kurien, R. Anuja, V. Hasyagar, A.P. Thomas,
R. Paliwal & J.M. Julka
(2023). Earthworm (Clitellata, Megadrili)
fauna of Kuttanad wetland, southern part of Vembanad-Kol Ramsar site, India. Opuscula
Zoologica, Budapest 54: 3–21.
Narayanan, M., J.M. Murugan, G. Kandasamy, S.
Kandasamy, O. Nasif, M. Rajendran & A. Pugazhendhi (2022). The
biotransformation potential of Bacillus cereus on β-cypermethrin to protect the
earthworm (Perionyx excavatus) on
insecticide-contaminated soil. Archives of Agronomy and Soil Science
68(7): 944–955.
Nesemann, H., G.
Sharma & R.K. Sinha (2004). Aquatic Annelida (Polychaeta, Oligochaeta, Hirudinea) of the Ganga
River and adjacent water bodies in Patna (India: Bihar), with description of a
new leech species (Family Salifidae). Annalen des Naturhistorischen
Museums in Wien, 105 B: 139–187. http://www.landesmuseum.at/pdf_frei_remote/ANNA_105B_0139-0187.pdf
Nesemann, H., S.
Sharma, G. Sharma, S.N. Khanal, B. Pradhan, D.N. Shah
& R.D. Tachamo (2007). Aquatic
Invertebrates of the Ganga River System, Volume 1 – Mollusca, Annelida,
Crustacea (in part). Chandi Press, Kathmandu, 263
pp.
Rao, C.R.N. (1922). Some new species of earthworms belonging to the
genus Glyphidrilus. Annals and Magazine of
Natural History, Series 9, 9(49): 51–68.
Reynolds, J.W. & M.J. Wetzel (2025). Nomenclatura Oligochaetologica –
A Catalogue of Names, Descriptions and Type Specimens of the Oligochaeta. Editio Secunda. https://nomenclatura-oligochaetologica.inhs.illinois.edu
. Accessed 20.ix.2025.
Reynolds, J.W., J.M. Julka & M.N. Khan
(1995). Additional earthworm records from Bangladesh (Oligochaeta:
Glossoscolecidae, Megascolecidae,
Moniligastridae, Ocnerodrilidae
and Octochaetidae). Megadrilogica
6(6): 51–60.
Rosa, D. (1890). Viaggio di Leonardo Fea in Birmania e regioni vicine, XXV. Moniligastridi, Geoscolecidi ed Eudrilidi. Annali
del Museo Civico di Storia Naturale di Genova,
Serie 2, 10: 107–122.
Stephenson, J. (1916). On a collection of Oligochaeta
belonging to the Indian Museum. Records of the Indian Museum 12: 99–354.
Stephenson, J. (1920). On a collection of Oligochaeta
from the lesser-known parts of India and from Eastern Persia. Memoirs of the
Indian Museum 7(3): 191–261.
Stephenson, J. (1921). Oligochaeta from
Manipur, the Laccadive Islands, Mysore and other parts of India. Records of
the Indian Museum 22: 745–768.
Stephenson, J. (1922). A note on some supposed new species of
earthworms of the genus Glyphidrilus.
Annals and Magazine of Natural History, Series 9, 9: 387–389. https://doi.org/10.1080/00222932208632688
Stephenson,
J. (1923). Oligochaeta. The Fauna of British India
including Ceylon and Burma. Taylor and Francis, London, xxiv + 518 pp.