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Biodiversity Data Journal :
Taxonomy & Inventories
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Corresponding author: Changfa Zhou (zhouchangfa@njnu.edu.cn)
Academic editor: Jean-Luc Gattolliat
Received: 14 Aug 2026 | Accepted: 18 Sep 2026 | Published: 29 Sep 2026
© 2026 Dewen Gong, Changfa Zhou
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gong D, Zhou C (2026) Reared adults of Potamanthellus unicutibius Nguyen & Bae, 2003 from China (Ephemeroptera, Neoephemeridae). Biodiversity Data Journal 14: e211811. https://doi.org/10.3897/BDJ.14.e211811
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The nymphs of Potamanthellus unicutibius Nguyen & Bae, 2003 possess filtering mouthparts and modified fore-legs. However, no adults are known.
In this study, both nymphal and imaginal stages of this species were collected and reared from China. Morphological examination reveals that these specimens differ markedly from all other members of Potamanthellus Lestage, 1931. In addition to the filtering nymphal morphology, the males of this species are characterised by more distinctly developed penes, longer tibiae and cerci and smaller compound eyes compared to those of Potamanthellus. Significantly, its distinct penes not only validates its independent species status, but also reveals the evolutionary process of the reduction of neoephemerid male genitalia and provides the original clue of the vestigial penes in several other lineages of the order Ephemeroptera as well.
evolution, imago, mayfly, Oriental realm, taxonomy
The species Potamanthellus unicutibius was originally described by
Unlike any other member of the family Neoephemeridae, nymphs of P. unicutibius possess filtering fore-legs, with both fore-tibiae and fore-tarsi bearing long, brush-like setae — a morphological trait typically observed in Isonychiidae or Oligoneuriidae (
Currently, the family Neoephemeridae comprises five extant genera and 17 described species (
In spring 2026, we conducted a collecting expedition in Yunnan Province, south-western China. During this survey, dozens of P. unicutibius nymphs were collected. Through careful indoor rearing, more than ten adults were successfully obtained. These newly-acquired materials provide a valuable opportunity to describe the previously unknown adult stage and to reassess the taxonomic and phylogenetic status of this enigmatic species.
Nymphs were collected using hand nets and adults were obtained both by rearing mature nymphs under laboratory conditions and by light trapping. All specimens were preserved in 80% or higher concentration ethanol and are currently deposited in the Mayfly Collection, College of Life Sciences, Nanjing Normal University (NNU), China.
For rearing, mature nymphs were placed in trays containing natural water, along with small stones, twigs and leaves. Each tray was covered with a small tent constructed from mosquito netting. Throughout the rearing period, a small aerator was used to continuously supply air to the water. Emerging adults were collected from inside the tent as they emerged.
Morphological observations were conducted using a combination of imaging techniques. Whole-body images of both nymphs and adults were captured with a Sony A7RIV interchangeable-lens digital camera, equipped with a 90 mm macro lens. For intermediate-sized structures, such as male genitalia and hind-wings, a Sony A7RIV camera, fitted with a Laowa FF 25 mm F2.8 Ultra Macro lens, was used. Egg morphology was examined using a Thermo Fisher Scientific Apreo 2S scanning electron microscope (SEM).
Eggs for SEM analysis were dissected from female imagoes. Together with the hind-legs of female subimagos, they were processed following a standard dehydration protocol: sequential immersion in an acetone gradient (70%, 80%, 90%, 98% and 100%, 10–20 min per step), followed by treatment with 1–1.5 ml of hexamethyldisilazane (HMDS) for 30 min. The specimens were then dried overnight in a vacuum desiccator and subsequently sputter-coated with a gold layer prior to imaging.
Potamanthellus unicutibius Nguyen & Bae, 2003 -
Potamanthellus unicutibius:
Male imago (in alcohol): body length 8.0–10.0 mm, fore-legs 11.0–12.0 mm, fore-wings 9.0–9.5 mm, hind-wings 3.0–3.2 mm, cerci 40.0–42.0 mm, terminal filament 1.0–1.1 mm; body reddish-brown (Fig.
Characters of male imago of Potamanthellus unicutibius Nguyen & Bae, 2003 A Head (dorsal view); B Fore-wing; C Hind-wing; D Hind-wing enlarged; E Fore-leg; F Fore-claw; G Mid-leg; H Mid-claw; I Hind-leg; J Genitalia (ventral view); K Genitalia (dorsal view); L Genitalia (apicodorsal view, showing penes).
Mouthparts and legs of nymph of Potamanthellus unicutibius Nguyen & Bae, 2003 A Labrum (dorsal view); B Left mandible; C Right mandible; D Hypopharynx (ventral view); E Maxilla; F Labium (ventral view); G Fore-leg; H Fore-tibia enlarged (arrows indicating the setaous rows); I Mid-leg; J Hind-leg
Nymphal characters of Potamanthellus unicutibius Nguyen & Bae, 2003 A Abdominal terga (arrows indicating the median tubercle); B Gills I; C Gills II (dorsal view, arrow indicating the ventral lamella); D Gills III (arrow indicating the ventral lamella); E Gills IV (arrow indicating the ventral lamella); F Gills VI; G Caudal filaments; H Cerci enlarged (showing the spines and setae).
Drawing pictures of Potamanthellus unicutibius Nguyen & Bae, 2003 A Nymphal habitus (dorsal view); B Fore-wing of male imago; C Hind-wing of male imago; D Male genitalia (ventral view, showing the forceps and subgenital plate); E Male genitalia (apicodorsal view, showing the penes and dorsal plate).
Female imago (in alcohol): body length 11.0–12.0 mm, fore-legs 5.0–6.0 mm, fore-wings 11.5–12.0 mm, hind-wings 3.5–4.5 mm, cerci 18.0–21.0, terminal filament 1.8–2.0 mm; body similar to male, but markings of wings slightly paler (Fig.
Male subimago (Fig.
Female subimago (Fig.
Mature nymph (in alcohol): (see
Main characters include: body length 8.0–12.0 mm, caudal filaments 3.0–4.0 mm. Body reddish-brown to chocolate (Fig.
Egg: surface smooth, micropyle at middle (Fig.
The nymph of P. unicutibius can be recognised by its filtering mouthparts and fore-legs: apical segment of maxillary and labial palpi more than 2 times of segment II in length, with dense hairlike setae (Fig.
In imaginal stage, its relatively short body (10.0–12.0 mm), heavily stained wings (Fig.
All other congeners in the genus Potamanthellus have no filtering setae on their fore-leg or mouthparts in nymphal stage. In imaginal stage, all other species have no distinct penes, only forceps and dorsal plate are visible. In addition, the dorsal plates of those species are different: P. amabilis and P. chinensis have deeply divided plates and their forceps are three-segmented. Although P. caenoides has two-segmented forceps, its dorsal plate is two semi-circular (see
Egg: surface smooth, without distinct mesh or KCTs (knob-terminated coiled threads), micropyle at equator (Fig.
Yunnan Prov. (China); Lao Cai Prov. (Vietnam).
Reduction or vestigialiation of male genitalia, particularly the penes, has been documented in several mayfly lineages. For instance, species within the taxon Cryptopenella of Choroterpes (Leptophlebiidae) possess indistinct penes, yet retain well-developed forceps and a distinct ventral penis cover (
In most mayflies, the gonopores open at the apex of the penes, implying that the penes are three-dimensional structures. However, based on historical accounts (
The genital morphology of P. unicutibius may reflect a possible evolutionary trend within the Neoephemeridae: initially, the forceps and penes are relatively well developed, as seen in Ephemera or Potamanthodes (
Within the Neoephemera-like clade, at least three nymphal morphotypes or evolutionary trends have been recognised: flattened body form (Pulchephemera), stout body form (Leucorhoenanthus and Neoephemera) and slender body form (Ochernova) (
From a biological perspective, based on general nymphal morphology, P. unicutibius and other Potamanthellus species appear to have evolved in different ecological directions. P. unicutibius may be adapted to filtering particulate matter in flowing water, whereas other Potamanthellus species likely exhibits greater swimming ability and may feed by chewing on fallen leaves or aquatic macrophytes.
This work was funded by the National Natural Science Foundation of China (Grant number 31750002, 32070475 and 32470464).
Dewen Gong collected materials of Potamanthellus unicutibius Nguyen & Bae, 2003.
Dewen Gong and Changfa Zhou made the illustrations, analysed the materials and wrote the descriptions and manuscript.