identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
039187C0FFD3FFF483CFFE7D37B364EA.text	039187C0FFD3FFF483CFFE7D37B364EA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon Bengtsson 1915	<div><p>Genus Procloeon Bengtsson 1915 (s. l.), or Procloeon/g1</p><p>(Figs 1–219)</p><p>In circumscription corresponds to:</p><p>—branch «11»: Kluge &amp; Novikova 1992: fig. 11;</p><p>— Procloeon /g1: Kluge 2012: 362;</p><p>—genus Procloeon: Kluge 2016: 494.</p><p>Type species: Cloeon bifidum Bengtsson 1912 .</p><p>Systematic position of Procloeon . The taxon Procloeon, or Procloeon /g1 belongs to the higher taxon Cloeon / fg1 (as defined by Kluge 2012), which can be treated as the tribe Cloeonini (as defined by Kluge 2016). Diagnosis and composition of Cloeon /fg1 = Cloeonini is given in the previous publications (Kluge &amp; Novikova 1992: branch «6» on figure 11; Kluge 2016: 492).</p><p>Diagnosis. The taxon Procloeon s. l. is characterized by peculiar structure of the denticles located on distal margins of segments of larval cerci: in distal part of cercus, each cercomere has one greatly enlarged denticle located on its lateral side; this denticle is much larger than others, usually longer than the next cercomere and is usually widened (Kluge &amp; Novikova 1992: fig. 9. 19) (Kluge 2012: 362; 2016: 494). Such structure of larval cerci occurs in all examined representatives if the plesiomorphon Procloeon s. str. (Figs 2–8; Kluge &amp; Novikova 1992: fig. 9. 19), the examined representative of the subgenus Pseudocentroptiloides s. str. (Fig. 1), in both representatives of the subgenus Monilistylus Kluge 2020a (Kluge et al. 2014: figs 1–3; Kluge 2020a: figs 7–8), in all representatives of the subgenus Oculogaster Kluge 2016 (Kluge 2016: figs 21–22, 59–60; Kluge 2020b: figs 8–9, 43–44, 85–86, 112, 141); in all examined representatives of the subgenus Securiops Jacobus et al. 2006 (Kluge 2023: figs 71–72, 105, 124–125) and in all examined representatives of the subgenus Psammonella described in this paper (Figs 86–88, 129–130, 156, 189). In most of these species, the enlarged denticle is longer than the next cercomere, in certain species up to three times longer (Kluge 2023: fig. 125); in exceptional case the denticle is a little shorter than the next cercomere (Kluge et al. 2014: fig. 3). The enlarged denticle is usually widened, with convex margins, so that in dorsal and ventral views its width at midlength exceeds its width at base (Figs 130, 156, 189); in some species these denticles are not widened, with margins straight (Fig. 86).</p><p>Psammophylous taxa within Procloeon</p><p>Larvae of some Procloeon s. l. are specialized for inhabitance on a bottom consisting mostly of clean sand. These are species of the subgenus Securiops (recently revised by Kluge, 2023) and the species originally placed by Jacob &amp; Glazaczov (1987) in the genus Pseudocentroptiloides .</p><p>These psammophylous larvae differ from other Procloeon by especially long and slender claws lacking denticles (Figs 74, 114–116, 151–153, 174–176) and by modifications of mouth parts. In contrast to other Procloeon, which have rather uniform mouth parts structure (Figs 9–17), the psammophylous larvae have labium enlarged and directed backward (Fig. 18), glossae diminished, paraglossae elongated, and labial palps especially greatly widened apically (Fig. 23; Kluge 2023: figs 40, 116).</p><p>Besides this, the genus Pseudocentroptiloides in the original sense (i.e., including the subgenus Psammonella) differs from Securiops and other Procloeon s. l. by widely incised distal margin of labrum (Fig. 19), widened and concave distal margin of hypopharynx (Fig. 22) and shortened maxillae with more or less developed setal crest laterad of incisors (Figs 24, 25–31). This lateral transverse setal crest originates from a row of setae which are few and small in other species (Fig. 194). These features of mouth parts structure are regarded to be diagnostic characters of the genus Pseudocentroptiloides s. l. (including the subgenus Psammonella) (Jacob &amp; Glazaczow 1987).</p><p>However, a recently described species Cheleocloeon psammonella Kluge 2025 demonstrates that similar shape of labrum, maxillae, glossae and paraglossae can independently evolve in a non-related species which has psammophylous specialization (Kluge 2025: figs 56, 59–63). This fact does not disprove the idea about holophyly of Pseudocentroptiloides s. l., but makes it less strongly argued.</p><p>Larva of Procloeon macronyx (Kluge &amp; Novikova 1992) has long and slender claws lacking denticles, like that of Pseudocentroptiloides; eggs of this species have equatorial girdle (Fig. 218) similar to that of Procloeon (Pseudocentroptiloides) shadini (Fig. 215); imagines of these two species are very similar. Both species are known under the name « Centroptilum nana » (see discussion below, «Status of species name Centroptilum nana »). Possibly, P. macronyx is related to P. shadini, but in contrast to it has no features of psammophylous specialization in structure of labrum, hypopharynx, maxillae and labium (Figs 9–17). Based on this, Belfiore &amp; D’Antonio 1990 placed both species in the genus Pseudocentroptilum, to which they attributed all Procloeon having hind wings, and regarded Pseudocentroptiloides to be a synonym of Pseudocentroptilum .</p><p>The new species, Procloeon furcalabrum sp. nov., described below, also has long and slender claws (Fig. 202); besides this, it has distal margin of labrum widely incised (Fig. 190) and apex of hypopharynx widened and concave (Fig. 195), as in Pseudocentroptiloides s. l.; but its maxillae are not shortened and retain long, slender canines (Fig. 194); its glossae and paraglossae retain the shape usual for Procloeon s. str. (Figs 191–192).</p><p>The equatorial girdle on eggs, being present in P. macronyx and P. (Pseudocentroptiloides) shadini, is absent in P. (Psammonella) magnificum sp. nov. (Figs 103–108), that suggests independent origin of Pseudocentroptiloides and Psammonella.</p><p>While status and circumscription of the taxa Pseudocentroptiloides s. l. (including Psammonella) and Pseudocentroptiloides s. str. (without Psammonella) are unclear, the taxon Psammonella represents a clearly holophyletic taxon comprising not less than 6 species distributed on Oriental Region. Holophyly of Psammonella is proven by unique structure of its maxillae (see below).</p></div>	https://treatment.plazi.org/id/039187C0FFD3FFF483CFFE7D37B364EA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFD5FFF483CFF9A837366758.text	039187C0FFD5FFF483CFF9A837366758.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Securiops) primasia (Kaltenbach 2023)	<div><p>Procloeon (Securiops) primasia (Kaltenbach et al. 2023)</p><p>Securiops primasia Kaltenbach, Phlai-ngam, Suttinun &amp; Gattolliat 2023: 129 (larva).</p><p>Procloeon (Securiops) primasia: Kluge 2023: 264 (larva, subimago, ♂ &amp; ♀ imago, eggs).</p><p>New finding. LAOS, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.915276&amp;materialsCitation.latitude=13.961666" title="Search Plazi for locations around (long 105.915276/lat 13.961666)">river Mekong</a> near lower end of island Don Det, 13°57'42"N, 105°54'55"E, 6–11.II.2025, coll. N. Kluge &amp; L. Sheyko: 2 L-S-I ♀, 2 larvae .</p></div>	https://treatment.plazi.org/id/039187C0FFD5FFF483CFF9A837366758	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFD5FFF483CFFB7137AE6591.text	039187C0FFD5FFF483CFFB7137AE6591.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Securiops Jacobus 2006	<div><p>Subgenus Securiops Jacobus et al. 2006</p><p>Genus Securiops Jacobus, McCafferty &amp; Gattolliat 2006: 133 (larva).</p><p>Subgenus Securiops Kluge 2020a: 577 (larva, male imago, egg).</p><p>Type species: Potamocloeon macafertiorum Lugo-Ortiz (in Lugo-Ortiz &amp; McCafferty) 1996.</p><p>Diagnosis and composition. See Kluge (2023).</p></div>	https://treatment.plazi.org/id/039187C0FFD5FFF483CFFB7137AE6591	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFD7FFF683CFFF1430D3678F.text	039187C0FFD7FFF683CFFF1430D3678F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	(Psammonella) Glazaczow 1987	<div><p>Subgenus Psammonella Glazaczow 1987</p><p>(Figs 97–189)</p><p>Psammonella Glazaczow (in Jacob &amp; Glazaczow) 1987: 203.</p><p>Type species: Pseudocentroptiloides (Psammonella) ceylonica Glazaczow (in Jacob &amp; Glazaczow) 1987.</p><p>Autapomorphy of Psammonella. Maxilla, being shortened (as in Pseudocentroptiloides s. str.), is apically armed with 3 transverse crests—two transverse crests formed by setae, and a shorter lamellate transverse crest between them (Figs 27–31).</p><p>The transverse setal crest laterad of canines (l.s.cr) represents a very dense and regular row of long, stout, hooked setae, which are projected distad of other two crests (Fig. 30); corresponding crest is present in Pseudocentroptiloides s. str. (Figs 25, 26); in other taxa setae corresponding to this crest are few and small (Fig. 194).</p><p>The transverse crest formed by fused canines (can.cr) is integral and plate-like, with convex lateral surface and concave median surface (Fig. 27); it corresponds to three separate canines, which are shortened in Pseudocentroptiloides s. str. (Figs 25–26) and are long in most other Procloeon (Fig. 194).</p><p>The transverse setal crest mediad of canines (m.s.cr) is a unique formation not found in other mayflies. It represents a distal continuation of the longitudinal medio-ventral row of setae which is sharply bent in dorsal direction, changing its orientation from longitudinal to transverse one (Figs 28–29). Setae forming this crest are longer than the plate-like crest formed by fused canines, and are curved S-sharply, so that are partly inserted into the concavity of the plate-like crest (Fig. 27). In all other mayflies the medio-ventral setal row, as well as the neighboring medio-dorsal setal row, is terminated close to canines, is not bent apically and does not form a transverse portion (Figs 25–26).</p><p>General characteristic of Psammonella. Labrum widened distally, with distal margin widely incised (Figs 67, 117, 144, 165) (the same in Pseudocentroptiloides and Procloeon furcalabrum sp. nov.). Incisor and kinetodontium of each mandible separated nearly up to base (Figs 72–73, 121–122, 145–146, 166–167) (the same in Pseudocentroptiloides, P. furcalabrum sp. nov. and P. macronyx). Maxillae modified (see above). Maxillary palp 2- or 3-segmented, or variable [as in other Procloeon (Figs 15–17)]. Hypopharynx apically widened and concave, with projection in middle of concavity, i.e. with 3 apices (Figs 71, 118, 149, 170). Glossae short and wide; paraglossae longer and narrower than glossae, arched, parallel-sided (Figs 69–70, 119, 147–148, 173). Hind wings of winged stages and hind protoptera of larvae completely absent in all known species (Figs 65, 113, 139) (the same in many other Procloeon). Claws long and slender, without denticles (Figs 74, 114–116, 151–153, 174–177). Tergalii either with dorsal lamella (Figs 46–49), or without it (Figs 32–45, 50–58).</p><p>Key to larvae of Psammonella</p><p>1(2) Tergalii I–IV with small dorsal lamella (Figs 46–49).................... Procloeon (Psammonella) diplofolium sp. nov.</p><p>2(1) All tergalii without dorsa lamella......................................................................... 3</p><p>3(4) Length of glossa less than width (Waltz &amp; McCafferty 1989: fig. 18)............... Procloeon (Psammonella) christinae</p><p>4(3) Length of glossa more than width (Figs 69, 119, 147–148, 173)................................................. 5</p><p>5(6) Outer side of femur with stout, spine-like setae (Fig. 77). Tergalius VII not stretched posteriorly (Fig. 38). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere not widened (Fig. 86)........................................................................................... Procloeon (Psammonella) magnificum sp. nov.</p><p>6(5) Outer side of femur without stout setae (Fig. 131). Tergalius VII stretched posteriorly (Figs 45, 59). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere widened (Figs 156, 189)................................. 7</p><p>7(8) Femur with dark brown cuticular band near middle (Fig. 177). Abdominal tergum and sternum IX with dark brown cuticular markings, terga III –VII without cuticular pigmentation (Figs 159–164). Tergalii maculated; tergalii I– VI roundish (Fig. 53– 59)........................................................... Procloeon (Psammonella) flavonigrum sp. nov.</p><p>8(7) Femur without dark cuticular band (only hypodermal band can be present— Figs 151–153). If abdominal terga have brown cuticular pigmentation, it is present at least on tergum VI (Figs 139, 142; Jacob &amp; Glazaczov 1986: fig. 8). Tergalii not maculated; tergalii II– V elongate (Figs 40–43; Jacob &amp; Glazaczov 1986: fig. 5).</p><p>9(10) 3rd segment of labial palp with inner angle sharply stretched (Figs 147–148)................................................................................................... Procloeon (Psammonella) mekongiense sp. nov.</p><p>10(9) 3rd segment of labial palp with inner angle not sharply stretched (Jacob &amp; Glazaczov 1986: fig. 3)........................................................................................... Procloeon (Psammonella) ceylonicum</p></div>	https://treatment.plazi.org/id/039187C0FFD7FFF683CFFF1430D3678F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFDBFFFE83CFFF14316D6781.text	039187C0FFDBFFFE83CFFF14316D6781.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Psammonella) magnificum Kluge 2025	<div><p>Procloeon (Psammonella) magnificum sp. nov.</p><p>(Figs 29–30, 32–38, 60–109)</p><p>Material examined. Holotype: L-S-I ♂ {specimen [XX](29)2020}, SRI LANKA, Sabaragamuwa Province, southeastern slope of Sri Pada, river Battulu oya, Fishing Hut, 9.II.2020, coll. N. Kluge &amp; L. Sheyko. Paratypes: the same locality and collectors, 8–10.II.2020, 1 L-S-I ♂, 1 L-S-I ♀, 1 L-S ♀, 90 larvae.</p><p>Etymology. The adjective magnificum (magnificent) is connected with the size larger that in other Psammonella, and with extensive reddish pigmentation of imago.</p><p>Larva. CUTICULAR COLORATION. Head from brown in dorsal (posterior) part to colorless in ventral (anterior) part (Fig. 61). Pronotum and mesonotum brown with diffusive maculation (Fig. 62). Metanotum from brown medially to colorless laterally (Fig. 65). Thoracic pleura and sterna colorless. Femur of each leg light ochre with outer side darker brown. Color of tibia and tarsus gradually changing from light ochre at proximal part of tibia to darker brown at distal part of tarsus (Figs 63–65, 74). Abdominal terga with darker brown and lighter ochre areas; terga II and VI darker than others (Fig. 60). Sterna colorless. Caudalii light ochre, with darker brownish margin of each 4th segment; swimming setae light ochre (Fig. 66).</p><p>HYPODERMAL COLORATION.Abdomen with or without transverse reddish bands on articulatory membranes between terga.</p><p>SHAPE AND SETATION. Labrum longer than in other species, with distal margin sharply concave (Figs 67– 68). Mandibles with incisor and kinetodontium separated up to base (Figs 72–73). Maxilla modified as characteristic for Psammonella (Figs 29–30, 71). Maxillary palp with 3rd (terminal) segment as long as 2nd segment (Fig. 71). Hypopharynx with 3 apices (Fig. 71). Glossae short and wide, obliquely truncated, laterally longer than medially; paraglossae longer and narrower than glossae, arched, either parallel-sided, or narrow at base and widened toward apex (Figs 69–70). 3rd (distal) segment of labial palp with inner angle not sharply stretched and distal margin straight (Figs 69–70).</p><p>Hind protoptera completely absent (Fig. 65).</p><p>Legs with thin, rather long, pointed, stout, two-channel setae; on femur such setae form longitudinal stripe on outer side, longitudinal stripe on inner side and longitudinal stripe on proximal part of posterior side (Figs 74, 76–77); on tibia such setae located on inner and posterior sides (Figs 78–79); on tarsus such setae form regular longitudinal row on inner side, few such setae present on posterior side (Figs 74–75). Claws long and slender, without denticles.</p><p>Abdominal segments II–VIII with pair of posterolateral denticles; segment IX with few denticles on lateral side near posterior margin. Posterior margin of abdominal tergum I without denticles; that of terga II–X with brown, pointed denticles alternated with smaller denticles; denticles of tergum II shortest, other longer (Fig. 81). Posterior margins of abdominal sterna I–II without denticles; that of sternum III with irregular denticles; that of sterna IV–IX with colorless, long, thin, sharply pointed denticles longer than denticles on terga (Fig. 82). Paraprocts with long and short, pointed denticles (Fig. 83). Each abdominal sternum II–VI with pair of sparse, transverse rows of long, colorless, bifurcate setae (Fig. 80).</p><p>Tergalii (Figs 32–38) without dorsal lamella; tergalii I–VI elongate; tergalius VII short and roundish, not stretched posteriorly (i.e. not of Procloeon - type).</p><p>Cerci and paracercus without sharply enlarged denticles on dorsal and ventral sides (Figs 84–86). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere (peculiar for Procloeon) thin, spine-like, 1.5 times longer than cercomere (Fig. 86).</p><p>POSE OF SUBIMAGINAL GONOSTYLI UNDER LARVAL CUTICLE of « Cloeon - type » (Fig. 94).</p><p>Subimago. CUTICULAR COLORATION. Head colorless, antennae brown. Pronotum ochre or light brownish. Mesonotum mostly light ochre, with margins of scutellum darker brown (Fig. 89). Sclerites of thorax (including postsubalar sclerite and lateropostnotal crest) ochre, as light as membranes and poorly visible (Fig. 90). Wing membrane colorless with microtrichia dark brown. Legs light ochre-brownish; apex of femur and patellar area of tibia darkened with brown. Abdominal terga nearly uniformly light ochre-brownish, sterna lighter.</p><p>HYPODERMAL COLORATION. As in imago.</p><p>TEXTURE. In both sexes, on all leg pairs, all tarsomeres are covered with pointed microlepides (Figs 98–99).</p><p>Imago, male (Figs 91–93). Head ochre-brown. Turbinate eyes yellow, with facetted surface widened in longitudinal direction. Thorax with ochre, brown and reddish areas. Fore wing colorless, with veins light ochre, with contrasting dark brown spot on costal brace. Hind wings absent. Legs ochre; apex of femur more or less darkened with reddish brown (as in Fig. 100). Middle and hind legs without apical spine on 1st+2nd tarsomere, with single apical spine on primary 3rd tarsomere (as in Fig. 98). Abdominal terga mostly ochre, with dark purple band on posterior margin; terga VIII–IX entirely dark purple; tergum X ochre with purple; lateral tracheal trunks dark gray. Sterna light ochre, only sternum VIII darker reddish. Gonostyli ochre. Penial bridge projected as wide, semicircular lobe (Figs 96–97).</p><p>Imago, female (Figs 101–102). Head ochre with brown. Coloration of thorax, wings and legs as in male. Abdominal terga reddish with ochre.</p><p>Egg (Figs 103–108). Oval. Surface entirely and evenly covered with fine, colorless lace consisting of flat disks and net of treads (Figs 105–106) raised above egg surface on brown spines (Figs 107–108).</p><p>Dimension. Fore wing length (and approximated body length) 7–8 mm.</p><p>Distribution. Sri Lanka. Larvae were abundant in the river Battulu oya (Fig. 109), but not found in the river Seethagangula running on another slope of the same mount Sri Pada, in spite of intensive collecting. This species was not found in other places of Sri Lanka as well.</p><p>Comparison. Larva of the new species Procloeon (Psammonella) magnificum sp. nov. well differs from Pr. (Ps.) ceylonicum (Glazaczow in Jacob &amp; Glazaczow 1987), Pr. (Ps.) diplofolium sp. nov., Pr. (Ps.) mekongiense sp. nov. and Pr. (Ps.) flavonigrum sp. nov. by tergalius VII, which is not stretched posteriorly and resembles tergalii of other pairs (Fig. 38). It well differs from these species also by presence of spine-like setae on outer side of femur (Fig. 77). It also differs at least from Pr. (Ps.) diplofolium sp. nov., Pr. (Ps.) mekongiense sp. nov. and Pr. (Ps.) flavonigrum sp. nov. by spine-like (i.e. not thickened) shape of the enlarged denticles on outer sides of cerci.</p><p>Larva of Pr. (Ps.) magnificum sp. nov. differs from incompletely described Pr. (Ps.) christinae (Waltz &amp; McCafferty 1989) at least by less shortened glossae, which have length exceeding width (Figs 69–70) (while glossae of P. christinae have length less than width— Waltz &amp; McCafferty 1989: fig. 18).</p></div>	https://treatment.plazi.org/id/039187C0FFDBFFFE83CFFF14316D6781	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFC5FFE883CFFB7931B0667C.text	039187C0FFC5FFE883CFFB7931B0667C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Psammonella) diplofolium Kluge 2025	<div><p>Procloeon (Psammonella) diplofolium sp. nov.</p><p>(Figs 27–28, 46–52, 110–135)</p><p>Material examined. Holotype: L-S ♀ {specimen [XIII] (6)B 2015}, THAILAND, prov. Mae-Hong-Son, Pai, river Pai upstream Tan Jet Ton village, 9.II.2015, coll. N. Kluge &amp; L. Sheyko. Paratypes: the same locality and collectors, 5–14.II.2015: 2 L-S ♀, 30 larvae .</p><p>Etymology. The species name diplofolium refers to double tergalii of some anterior pairs (Figs 46–49). While this character occurs in many other representatives of the genus Procloeon, it is unknown for other representatives of the subgenus Psammonella .</p><p>Larva. CUTICULAR COLORATION. Head mostly colorless, only frons dorsad of antennae bases and occipit colored with light ochre (Fig. 111). Pronotum and mesonotum ochre, with small, contrasting darker brown maculae (Fig. 112). Metanotum and thoracic pleura with brown and colorless areas (Fig. 113). Legs mostly colorless (Figs 114–116). Abdominal terga with darker brown and lighter ochre areas; terga I, III, V–VI and IX darker than others (Fig. 110). Sterna colorless. Caudalii light ochre, with darker brownish margin of each 4th segment; swimming setae with light bases, darker brownish middle parts and colorless tips (Fig. 110).</p><p>HYPODERMAL COLORATION. Abdominal terga with transverse reddish band at middle part of posterior margin; these or that abdominal terga with or without other unpaired and/or paired reddish markings (Fig. 134).</p><p>SHAPE AND SETATION. Labrum widened distally, with distal margin sharply concave (Fig. 117). Mandibles with incisor and kinetodontium separated up to base (Figs 121–122). Maxilla modified as characteristic for Psammonella (Figs 27–28, 120). Maxillary palp with 3rd (terminal) segment as long as 2nd segment (Fig. 120). Hypopharynx with 3 apices (Fig. 118). Glossae short and wide, rounded apically; paraglossae longer and narrower than glossae, arched, parallel-sided (Fig. 119). 3rd (distal) segment of labial palp with inner angle stretched and distal margin concave (Fig. 119).</p><p>Hind protoptera completely absent (Fig. 113). Legs with thin, rather long, pointed, stout, two-channel setae located on inner sides of femur, tibia and tarsus; no stout setae on outer side of leg (Fig. 131). Claws long and slender, without denticles (Figs 114–116.</p><p>Abdominal segments II–VII with pair of posterolateral denticles; segments VII–IX with several denticles on lateral sides. Posterior margin of abdominal tergum I with small, irregular denticles; that of terga II–X with pointed denticles alternated with small denticles or blunt protuberances (Fig. 123, 128). Posterior margins of abdominal sterna I–III without denticles; that of sterna IV–IX with pointed denticles shorter than denticles on terga (Fig. 124). Paraprocts with long, pointed denticles (Fig. 125). Sterna II–VIII with one or few colorless, bifurcate setae sublaterally (as on Fig. 182).</p><p>Tergalii I–IV with small dorsal lamella; tergalius VII stretched posteriorly, of usual Procloeon - type (Figs 46– 52).</p><p>In middle part of cerci and paracercus, denticles on posterior margin of each 4th segment equally enlarged on dorsal and ventral sides (Figs 126–127). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere (peculiar for Procloeon) widened at midlength and slightly longer than cercomere (Figs 129–130).</p><p>POSE OF SUBIMAGINAL GONOSTYLI UNDER LARVAL CUTICLE of « Cloeon - type » (as in Fig. 94).</p><p>Subimago (based on reared females and males extracted from larva). CUTICULAR COLORATION. Subimaginal cuticle mostly colorless; mesonotum with pair of brown stripes along lateroparapsidal sutures (Fig. 132). Wing membrane entirely colorless, microtrichia ochre.</p><p>HYPODERMAL COLORATION. Body ochre; all or some abdominal terga with more or less expressed transverse reddish band at middle part of posterior margin, sometimes with other reddish markings (Figs 134–135). Legs uniformly ochre (Fig. 133).</p><p>TEXTURE. In both sexes, on all leg pairs, all tarsomeres are covered with pointed microlepides (as in Figs 98–99) (Kluge 2022).</p><p>Imago. Unknown.</p><p>Eggs. Unknown. Eggs extracted from female subimagines are undeveloped.</p><p>Comparison. Larva of the new species Procloeon (Psammonella) diplofolium sp. nov. differs from all other known Psammonella by presence of dorsal lamella on tergalii of some pairs and by rounded glossae.</p></div>	https://treatment.plazi.org/id/039187C0FFC5FFE883CFFB7931B0667C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFC9FFED83CFF90A37CE648E.text	039187C0FFC9FFED83CFF90A37CE648E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Psammonella) mekongiense Kluge 2025	<div><p>Procloeon (Psammonella) mekongiense sp. nov.</p><p>(Figs 39–45, 136–158)</p><p>Material examined. Holotype: female larva ready to molt to subimago, LAOS, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.915276&amp;materialsCitation.latitude=13.961666" title="Search Plazi for locations around (long 105.915276/lat 13.961666)">river Mekong</a> near lower end of island Don Det, 13°57'42"N, 105°54'55"E, 6–11.II.2025, coll. N. Kluge &amp; L. Sheyko. Paratype: the same locality, date and collectors: 1 female larva of penultimate instar .</p><p>Etymology. The species name mekongiense refers to its inhabitance in river Mekong.</p><p>Larva. CUTICULAR COLORATION. Head mostly colorless, occipit with brownish maculae (Fig. 138). Pronotum and mesonotum with brown, ochre and colorless areas (Fig. 137). Metanotum and thoracic pleura with brown and colorless areas. Cuticle of legs mostly colorless. Abdominal terga with darker brown, lighter ochre and colorless areas; terga I, III, V–VI and IX darker than others (Fig. 139). Sterna colorless. Caudalii light ochre, with darker brownish margin of each 4th segment; swimming setae from ochre to colorless (Fig. 139).</p><p>HYPODERMAL COLORATION. Each abdominal tergum I–IX with unpaired longitudinal reddish stripe and pair of transverse reddish stripes near anterior margin; each tergum II and VIII with pair of reddish spots near lateral margins (Fig. 140).</p><p>SHAPE AND SETATION. Labrum widened distally, with distal margin sharply concave (Fig. 144). Mandibles with incisor and kinetodontium separated up to base (Figs 145–146). Maxilla modified as characteristic for Psammonella (Fig. 150). Maxillary palp with 3rd (terminal) segment shorter than 2nd segment (Fig. 150). Hypopharynx with 3 apices (Fig. 149). Glossae short and wide, with rounded margins, laterally longer than medially; paraglossae longer and narrower than glossae, arched, parallel-sided (Figs 147–148). 3rd (distal) segment of labial palp with inner angle stretched and distal margin concave (Figs 147–148).</p><p>Hind protoptera completely absent (Fig. 139).</p><p>Legs with thin, rather long, pointed, stout, two-channel setae located on inner sides of femur, tibia and tarsus; no stout setae on outer side of leg (as in Figs 131). Claws long and slender, without denticles (Figs 151–153).</p><p>Abdominal segments II–VII with pair of posterolateral denticles; segments VII–IX with several denticles on lateral sides (Fig. 155). Posterior margin of abdominal tergum I without denticles; that of terga II–X with pointed denticles alternated with small denticles or blunt protuberances (Fig. 155). Posterior margins of abdominal sterna I–III without denticles; that of sterna IV–IX with pointed denticles shorter than denticles on terga (Fig. 155). Paraprocts with long, pointed denticles (Fig. 155). [Bifurcate setae on sterna not examined]</p><p>Tergalii (Figs 39–45) without dorsal lamella; tergalii I– V elongate; tergalius VI short and roundish; tergalius VII stretched posteriorly, of usual Procloeon - type (Fig. 45).</p><p>In middle part of cerci and paracercus, denticles on posterior margin of each 4th segment nearly equally enlarged on dorsal and ventral sides (Figs 157–158). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere (peculiar for Procloeon) widened at midlength and slightly longer than cercomere (Fig. 156).</p><p>Subimago (based on female subimago extracted from larva). CUTICULAR COLORATION. Cuticle very light or colorless; mesonotum colorless, only anterolateral scutal crest and prelateroscutum darkened with brownish (Fig. 154). Wing membrane entirely colorless, microtrichia ochre.</p><p>HYPODERMAL COLORATION. Wings crumpled under larval cuticle in proximal part dark gray, in distal part yellowish (Fig. 154). Legs of female with trochanter, femur, tibia and tarsus mostly light ochre, femur darkened with brown near base and near apex, tibia darkened with brown at outer side (Figs 151–153). Each abdominal tergum I–IX with unpaired longitudinal reddish stripe and pair of transverse reddish stripes near anterior margin; each tergum II and VIII with pair of reddish spots near lateral margins (Figs 136, 140, 142–143).</p><p>TEXTURE. In both sexes, on all leg pairs, all tarsomeres are covered with pointed microlepides (as in Figs 98–99) (Kluge 2022).</p><p>Imago. Unknown.</p><p>Eggs. Unknown. Eggs extracted from mature female larva are undeveloped.</p><p>Comparison. Larva of the new species Procloeon (Psammonella) mekongiense sp. nov. differs from Pr. (Ps.) ceylonicum (Glazaczow in Jacob &amp; Glazaczow 1987) by stretched inner angle of 3rd segment of labial palp. It differs from incompletely described Pr. (Ps.) christinae (Waltz &amp; McCafferty 1989) at least by less shortened glossae, which have length exceeding width (Figs 147–148) (while glossae of P. christinae have length less than width— Waltz &amp; McCafferty 1989: fig. 18).</p></div>	https://treatment.plazi.org/id/039187C0FFC9FFED83CFF90A37CE648E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFCCFFEE83CFFA9531D066AA.text	039187C0FFCCFFEE83CFFA9531D066AA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Psammonella) flavonigrum Kluge 2025	<div><p>Procloeon (Psammonella) flavonigrum sp. nov.</p><p>(Figs 31, 53–59, 159–189)</p><p>Material examined. Holotype: male larva ready to molt to subimago, INDIA, state Kerala, river Periyar in Malaytoor, 15.II.2015, coll. N. Kluge &amp; L. Sheyko. Paratypes: the same locality, date and collectors: 6 larvae .</p><p>Etymology. The species name flavonigrum refers to bright yellowish color of larva with contrasting black markings on metanotum, abdomen and femora (Fig 159).</p><p>Larva. CUTICULAR COLORATION. Head light ochre. Pronotum and mesonotum with protoptera light ochre, tips of longitudinal veins darkened with brownish (Fig. 179). Cuticle of leg from light ochre to colorless; each femur with narrow, contrasting, dark brown band distad of midlength (Figs 159, 161, 174–177). Cuticle of abdomen mostly colorless, with very contrasting dark brown (nearly black) markings: in most pigmented individuals, dark brown are most part of metanotum, most part of abdominal tergum I, maculae on tergum II, unpaired macula on tergum VIII, most part of tergum IX, small macula on tergum X, pair of maculae near anterior margin of sternum VIII and most part of sternum IX (Figs 159–164). In two individuals dark brown cuticular pigmentation of abdomen is limited by pair of lateral lines on tergum IX and paired markings on sternum IX (Figs 161–162). Caudalii light ochre, with darker brownish joining of each 4th segment; swimming setae with light bases, darker brownish middle parts and colorless tips (Fig. 186).</p><p>HYPODERMAL COLORATION. These or that abdominal terga with or without unpaired and/or paired reddish markings (Fig. 161). Tergalii maculated due to dark brown pigmentation on some areas of tracheae (Figs 53–59).</p><p>SHAPE AND SETATION. Labrum widened distally, with distal margin sharply concave (Fig. 165). Mandibles with incisor and kinetodontium separated up to base (Figs 166–167). Maxilla modified as characteristic for Psammonella (Figs 31, 171). Maxillary palp with 3rd (terminal) segment shorter than 2nd segment (Fig. 171). Hypopharynx with 3 apices (Fig. 170). Glossae short and wide, with rounded margins, laterally longer than medially; paraglossae longer and narrower than glossae, arched, parallel-sided (Fig. 168, 173). 3rd (distal) segment of labial palp with inner angle stretched and distal margin concave (Fig. 168–172); stretching and concavity not expressed on left palp of holotype (right side of Fig. 168).</p><p>Hind protoptera completely absent (Fig. 164).</p><p>Legs with thin, rather long, pointed, stout, two-channel setae located on inner sides of trochanter, femur, tibia and tarsus; no stout setae on outer side of leg (Fig. 178). Claws long and slender, without denticles (Figs 174–176).</p><p>Abdominal segments II–VII with pair of posterolateral denticles; segments VIII–IX with several denticles on lateral sides. Posterior margin of abdominal tergum I without denticles; that of terga II–X with pointed denticles alternated with small denticles or blunt protuberances (Fig. 181, 184). Posterior margins of abdominal sterna I–III without denticles; that of sterna IV–V with or without denticles; that of sterna VI–IX with pointed denticles shorter than denticles on terga (Fig. 183). Paraprocts with long, pointed denticles (Fig. 185). Sterna II–VIII with one or few colorless, bifurcate setae sublaterally (Fig. 182).</p><p>Tergalii (Figs 53–59) without dorsal lamella; tergalii I–VI short and roundish; tergalius VII stretched posteriorly, of usual Procloeon - type (Fig. 59). Tracheae of tergalii partly clean and colorless, partly densely covered with pigmented granules, that makes tergalii maculated.</p><p>In middle part of cerci and paracercus, denticles on posterior margin of each 4th segment enlarged on dorsal side (Fig. 187) and smaller on ventral side (Fig. 188). In distal part of cercus, greatly enlarged denticle on lateral side of each cercomere (peculiar for Procloeon) widened at midlength and slightly longer than cercomere (Fig. 189).</p><p>POSE OF SUBIMAGINAL GONOSTYLI UNDER LARVAL CUTICLE of « Cloeon - type » (as in Fig. 94).</p><p>Subimago (based on male subimago extracted from larva).</p><p>CUTICULAR COLORATION. Cuticle very light or colorless; mesonotum colorless. Wing membrane entirely colorless, microtrichia brown.</p><p>HYPODERMAL COLORATION. Wings crumpled under larval cuticle in proximal part light ochre, in middle part darker brow, in distal part yellowish; large, dark, purple-brown macula near costal brace (Fig. 180).</p><p>Legs with trochanter dark brown; tibia and tarsus ochre, tibia darkened with brown near base and indistinctly darkened near apex; femur of fore leg reddish-brown (Fig. 174); femora of middle and hind legs ochre, darkened with reddish-brown near apex (Figs 175–176). These or that abdominal terga with or without unpaired and/or paired reddish markings (Figs 161, 164).</p><p>TEXTURE. In both sexes, on all leg pairs, all tarsomeres are covered with pointed microlepides (Figs 98–99) (Kluge 2022).</p><p>Imago. Unknown.</p><p>Eggs. Unknown.</p><p>Dimension. Body length of last instar larva 4 mm.</p><p>Comparison. Larva of the new species Procloeon (Psammonella) flavonigrum sp. nov. differs from Procloeon (Psammonella) ceylonicum (Glazaczow in Jacob &amp; Glazaczow 1987) by stretched inner angle of 3rd segment of labial palp, dark brown band near middle of each femur, dark brown markings on abdominal tergum and sternum IX in line with absence of pigmentation on terga III–VII, and by roundish, maculated tergalii.</p><p>Larva of Pr. (Ps.) flavonigrum sp. nov. differs from incompletely described Pr. (Ps.) christinae (Waltz &amp; McCafferty 1989) at least by less shortened glossae, which have length exceeding width (Fig.) (while glossae of P. christinae have length less than width— Waltz &amp; McCafferty 1989: fig. 18) and by shorter, roundish tergalius IV (Fig. 57) (while in P. christinae this tergalius is longer— Waltz &amp; McCafferty 1989: fig. 23).</p></div>	https://treatment.plazi.org/id/039187C0FFCCFFEE83CFFA9531D066AA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFF2FFD383CFFA6C31AC676F.text	039187C0FFF2FFD383CFFA6C31AC676F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	(Pseudocentroptiloides) Jacob 1987	<div><p>Subgenus Pseudocentroptiloides Jacob 1987</p><p>(Figs 1, 18–24, 25–26, 207–209, 213–216)</p><p>Subgenus Pseudocentroptiloides Jacob (in Jacob &amp; Glazaczow) 1987: 203 (larva).</p><p>Type species: Pseudocentroptilum shadini Kazlauskas 1964 .</p><p>Diagnosis. See Jacob &amp; Glazaczow (1987).</p><p>Composition. Palaearctic species Procloeon (Pseudocentroptiloides) shadini (Kazlauskas 1964) and two Nearctic species, P. usa Waltz &amp; McCafferty 1989 and P. morihari Wiersema &amp; McCafferty 1998 .</p></div>	https://treatment.plazi.org/id/039187C0FFF2FFD383CFFA6C31AC676F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFF2FFD383CFFF5C37C165D5.text	039187C0FFF2FFD383CFFF5C37C165D5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (subgen.?) Bengtsson 1915	<div><p>Procloeon (subgen.?) furcalabrum sp. nov.</p><p>(Figs 190–203)</p><p>Material examined. Holotype: female larva of penultimate instar, THAILAND, prov. Mae-Hong-Son, Pai, river Pai upstream Tan Jet Ton village, 5–11.II.2015, coll. N. Kluge &amp; L. Sheyko.</p><p>Etymology. The species name furcalabrum refers to bifurcate shape of labrum due to its concave distal margin (Fig. 190); this character is common with Psammonella and Pseudocentroptilum, from which the new species differs by non-modified maxillae and labium.</p><p>Larva. CUTICULAR COLORATION. Legs mostly ochre, each femur with brown macula in distal part (Fig. 203). Abdominal terga mostly ochre with brown markings: tergum II with lateral and middle markings, terga III and V with lateral markings, tergum VI mostly brown (Fig. 198).</p><p>HYPODERMAL COLORATION. Abdominal terga with paired, transverse, reddish spots at segment joinings; tergum I with pair of roundish, reddish spots laterally (Fig. 198).</p><p>SHAPE AND SETATION. Labrum with distal margin concave (Fig. 190). Mandibles with incisor and kinetodontium separated up to base (Figs 196–197). Maxilla with all 3 canines long, slender and pointed, with 3 long and slender dentisetae; inner-ventral and inner-dorsal setal rows not curved, setae laterad of canines few and not forming transverse row (Fig. 194). Maxillary palp 3-segmented (Fig. 193). Glossae and paraglossae pointed, of subequal length; 3rd (terminal) segment of labial palp moderately widened distally (Figs 191–192).</p><p>Hind protoptera completely absent.</p><p>Femur, tibia and tarsus without stout setae on outer side; inner side of femur with few small, stout, pointed setae; inner side of tibia and tarsus with longer, pointed setae. Claws long and slender, with 2 rows of small, slender denticles on proximal 1/3 of claw length (Fig. 202).</p><p>Abdominal segments II–VII with pair of posterolateral denticles; segments VIII–IX with several denticles on lateral sides (Fig. 200). Posterior margin of abdominal tergum I without denticles; that of terga II–X with pointed denticles alternated with small denticles or blunt protuberances (Figs 200–201). Posterior margins of abdominal sterna I–V without denticles; that of sterna VI–IX with pointed denticles shorter than denticles on terga (Fig. 199). Paraprocts with pointed denticles of various sizes (Fig. 199).</p><p>Tergalii without dorsal lamella; at least tergalii I and III pointed (Figs 204–206).</p><p>Imago and subimago. Unknown.</p><p>Eggs. Unknown.</p><p>Dimension. Body length of penultimate instar larva 5.5 mm.</p><p>Comparison. The new species Procloeon furcalabrum sp. nov. is a single known species of Procloeon which have concave distal margin of labrum, and at the same time has no modifications of labium and maxillae peculiar for Pseudocentroptiloides and Psammonella.</p></div>	https://treatment.plazi.org/id/039187C0FFF2FFD383CFFF5C37C165D5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFF7FFD683CFFDF53494668C.text	039187C0FFF7FFD683CFFDF53494668C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Procloeon (Pseudocentroptiloides) shadini (Kazlauskas 1964)	<div><p>Procloeon (Pseudocentroptiloides) shadini (Kazlauskas 1964)</p><p>(Figs 1, 18–24, 25–26, 207–209, 213–216)</p><p>Baetidae gen. sp.: Neizvestnova-Zhadina 1931: 167 (lava).</p><p>Pseudocentroptilum? shadini Kazlauskas 1964: 170 (larva); Kluge 1995: 15 (lectotype designation).</p><p>Centroptilum shadini: Keffermüller 1978: 99 (♀ imago); Keffermüller &amp; Sowa 1984: 327 (♂ &amp; ♀ imago, larva).</p><p>Pseudocentroptiloides (Pseudocentroptiloides) shadini: Jacob &amp; Glazaczow 1987: 203 (larva).</p><p>Pseudocentroptiloides shadini: Waltz &amp; McCafferty 1989: 154 (larva, ♂ imago); Glazaczow &amp; Klonowska-Olejnik 2009: 151 (larva, ♂ &amp; ♀ imago, egg).</p><p>Cloeon (Pseudocentroptiloides) nana: Kluge &amp; Novikova 1992: 80 (subimago, ♂ &amp; ♀ imago, egg).</p><p>Descriptions. See Keffermüller &amp; Sowa (1984); Kluge &amp; Novikova (1992).</p><p>Distribution. Palaearctic. Reported from France (Chovet &amp; Brulin 2016), Poland (Keffermüller &amp; Sowa 1984; Glazaczow &amp; Klonowska-Olejnik 2009), Turkey (Kazanci 2001), Central Russia (Kazlauskas 1964) and Northern Urals (Novikova 1984). Reared specimens examined by the author were collected in Lithuania and Urals; besides this, there are imagines (but not larvae) from Evenkia in Eastern Siberia (Kluge &amp; Novikova 1992).</p><p>Disagreement about structure of tergalii. For the first time, 15 larvae of this species were reported from river Oka by Neizvestnova-Zhadina (1931) under the name « Baetidae gen. sp.». Neizvestnova-Zhadina (1931: fig. 2) gave drawings of maxilla, labium and tergalius of the 2nd pair which was the single preserved tergalius among all examined individuals. The maxilla and the labium undoubtedly belong to P. shadini . The tergalius has the 2nd (dorsal) lamella (Neizvestnova-Zhadina 1931: fig. 2c).</p><p>Among 8 larvae originally described by Kazlauskas (1964) as Pseudocentroptilum s hadini, only few ones had tergalius of the 1st pair only, and these tergalii had no dorsal lamella. Comparing these facts, Kazlauskas (1964) assumed that in P. s hadini tergalii of the 1st pair lack dorsal lamella, and the next tergalii have it.</p><p>Subsequent examinations of complete individuals showed that this conclusion was wrong, and actually all tergalii of this species lack dorsal lamella (Keffermüller &amp; Sowa 1984: 330, fig. 27).</p><p>Possibly, this disagreement was caused by the fact that Neizvestnova-Zhadina confused larvae of P. shadini with Procloeon macronyx . Kazlauskas (1964) reported larvae presumably determined by him as « Centroptilum nana Bogoescu » from the river Oka; according to his description, these larvae belong to P. macronyx . Neizvestnova-Zhadina (1931) did not report larvae of P. macronyx from the river Oka. It is not excluded, that she confused them with the larvae of P. shadini, and draw the maxilla and the labium taken from P. shadini, and the tergalius of 2nd pair taken from P. macronyx .</p></div>	https://treatment.plazi.org/id/039187C0FFF7FFD683CFFDF53494668C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFF6FFD783CFFC9D301766F6.text	039187C0FFF6FFD783CFFC9D301766F6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Centroptilum romanicum Bogoescu 1949	<div><p>Status of the species name Centroptilum romanicum Bogoescu 1949</p><p>The species under the name « Centroptilum romanicum » was described by Bogoescu (1949) based on male and female imagines and subimago collected in region Săcel (Gorj) in Romania. A list of specimens examined was not given, holotype was not designated, and place of material deposition was not reported. The type specimens are probably lost. Assumption that the male and the female belong to one and the same species, was based on nothing.</p><p>Identity of male imago. Figure of male genitalia (Bogoescu 1949: fig. 5) suggests that the genitalia were deformed due to muscle contraction, so the real shape of gonostyli and unistyligers is unknown. Subsequently, this figure was reproduced in some publications (Bogoescu 1958: fig. 76B; Keffermüller &amp; Sowa 1984: fig. 34; Bauernfeind &amp; Soldán 2012: fig. 117), but nobody redescribed male imago of this species based on original material.</p><p>Identity of female imago. In the original description (Bogoescu 1949), egg structure was not described. Eggs ascribed to Centroptilum romanicum were described later as following: «Ouăle au corionul prevăzut cu un brîu reticulat mai îngroşat, format din 13 rînduri de ochiuri (fig. 76, D)» (The eggs have a chorion provided with a thicker reticulated girdle, made up of 13 rows of meshes) (Bogoescu 1958: 126). According to this description and the drawing (Bogoescu 1958: fig. 76D), the egg structure completely agrees with that of P. shadini (Figs 213–216).</p><p>In another publication (Bogoescu &amp; Tabacaru 1966), Centroptilum romanicum was included in a key for female imagines, and its egg was characterized as «Die Äquatorialzone des Chorions mit 13–15 Warzenreihen» (the equatorial zone of chorion with 13–15 rows of warts). This wordily characteristics also agrees with P. shadini; but the drawing (Bogoescu &amp; Tabacaru 1966: fig. 3B) is different from the previous one, with hemispheric warts instead of papillae with apical concavities forming the reticulate relief. Material examined was not reported.</p><p>Interpretations of romanicum [ Centroptilum]. Under the name « Centroptilum romanicum », Keffermüller &amp; Sowa (1984) described a larva of P. macronyx with relatively large dorsal lamellae of tergalii and relatively short claws. This description was based on a single larval specimen from Romania. They also described eggs differently from the eggs described by Bogoescu: also with 13–15 rows of enlarged papillae, but with “single small papillae” in meshes of the dense net-like relief covering the rest surface (Keffermüller &amp; Sowa 1984: p. 331 and fig. 36).</p></div>	https://treatment.plazi.org/id/039187C0FFF6FFD783CFFC9D301766F6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
039187C0FFF6FFDA83CFF88D37E366AA.text	039187C0FFF6FFDA83CFF88D37E366AA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Centroptilum nana Bogoescu 1951	<div><p>Status of the species name Centroptilum nana Bogoescu 1951</p><p>The species under the name « Centroptilum nana » was described by Bogoescu (1951) based on male and female imagines collected in Ţugureşti (Dolj) near bank of Jiului in Romania. A list of specimens examined was not given, holotype was not designated, and place of material deposition was not reported. The type specimens are probably lost (Glazaczow &amp; Klonowska-Olejnik 2009: 150). Assumption that the male and the female belong to one and the same species, was based on nothing.</p><p>Species name. Judging by the fact that earlier this author introduced the binomen Centroptilum romanicum Bogoescu 1949 with the species name « romanicum » as adjective neutral with ending «- um », the species name « nana » is not an adjective, but a noun with non-changeable ending. In spite of this, some authors adopted it as adjective neutral—either as « Centroptilum nanum » (Keffermüller 1967), or « Cloeoptilum nanum » (Kazlauskas 1972), or « Procloeon nanum » (Jacob 1991), or « Pseudocentroptilum nanum » (Kovacs et al. 1998) .</p><p>Identity of male imago. Figure of male genitalia (Bogoescu 1951: fig. 2) suggests that the male imago originally described as Centroptilum nana, is conspecific either with Procloeon (Pseudocentroptiloides) shadini, or with Procloeon macronyx . Male imagines of these two species have no reliable difference in structure of genitalia and hind wings (Kluge &amp; Novikova 1992: figs 1, 2, 5, 11, 12, 14; Glazaczow &amp; Klonowska-Olejnik 2009: figs 29– 30, 36–37), but can be distinguished by coloration of abdomen. In the both male imaginal individuals of P. shadini reared by me from larvae collected in the river Neris in Lithuania, coloration of abdominal terga is differentiated as the following: terga II– III and V – VI have red-brown markings, which are absent on tergum IV (Figs 207–208; Kluge &amp; Novikova 1992: fig. 13 and p. 80). Among three male individuals reared by Keffermüller from larvae collected in river Warta in Poland, one individual has the same differentiation of abdominal terga: «po bokach, wzdłuż tylnej krawędzi oraz w środku segmentów (zwłaszcza II, III, V i VI) rdzawy nalot, wyraźny tylko u jednego z okazów» (Keffermüller 1967: 18). In contrast to this, all 15 male imagines of P. macronyx reared by me from larvae collected in the river Neris in Lithuania, had uniform red-brown markings on abdominal terga II– VI; tergum IV is not lighter than others, but, vice verse, with a pair of darker lateral markings (Fig. 211; Kluge &amp; Novikova 1992: fig. 3 and p. 77). All 15 male imagines of P. macronyx reared by me from larvae collected in the river Chu in Kazakhstan, also had uniform markings on abdominal terga II– VI, but less extensive than in the specimens from Lithuania (Fig. 210). After preservation in alcohol since 1986 and 1988 till 2025, these specimens lost their red-brown markings of abdominal terga; the coloration is preserved in the specimens mounted on slides in Canadian balsam.</p><p>According to the original description of Centroptilum nana, «Abdomenul are tergitele 1, 2, 7 si 8 pigmentate mult in brun inchis, tergitele 3–6 colorate foarte slab, iar 9 si 10 de coloare galben-brun» (The abdomen has tergites 1, 2, 7 and 8 highly pigmented in dark brown, tergites 3–6 very weakly colored, and 9 and 10 yellow-brown) (Bogoescu 1951: 2). In Baetidae, the first abdominal tergum of male imago is mostly substituted by metanotum, so terga «1, 2» could be confused with terga II and III. We (Kluge &amp; Novikova 1992) synonymized P. shadini with P. nana based on this assumption and on the fact that abdominal terga have differentiated color pattern in P. shadini and uniform color pattern in P. maronyx .</p><p>Identity of female imago. Eggs extracted from female imago of Centroptilum nana, were characterized as «Oul arc corionul înconjurat de un brâu reticulat ingrosat, alcătuit din 7 rânduri» (The egg chorion is surrounded by a thickened reticulate girdle, made up of 7 rows), and illustrated by a drawing (Bogoescu 1951: 3, fig. 4). According to this drawing, the egg is ellipsoid («spindle-shape» according to Glazaczow &amp; Klonowska-Olejnik 2009), i.e. gradually narrowing toward poles, and its girdle consists of cells projected above other cells of the egg surface. In another publication by this author, optic section of egg chorion is shown (Bogoescu 1958: fig. 77C), which demonstrates that inner margin of the girdle is straight, and its outer margin is convex. Such egg structure occurs in P. shadini (Figs 215–216).</p><p>In contrast to this, egg of P. macronyx (Figs 218–219) has hemispheric polar areas («oval shaped» according to Glazaczow &amp; Klonowska-Olejnik 2009) and a concave equatorial area, with a girdle inserted into this concavity; papillae of the girdle are terminated not by concavities, but by convexities, so they do not form cells of a reticulum, but form hemispheric projections located in the same plane as the cells of other egg surface (Fig. 217). In optic section, inner margin of the girdle is concave, and its outer margin is straight (Fig. 219).</p><p>Number of rows forming the girdle is reported and figured as 7 for Centroptilum nana . Eggs of P. macronyx examined by me have 5–11 rows, and eggs of P. shadini examined by me have 11–15 rows. Based on the rows number only, Glazaczow &amp; Klonowska-Olejnik (2009), concluded that Centroptilum nana is conspecific with P. macronyx . Concerning other features of the egg structure, they wrote: «It is difficult to state exactly what was seen by Bogoescu through the microscopes from the middle of the last century». However, it is clear that Bogoescu used translucent light microscopy and saw optic sections of eggs, as well as surfaces of eggs, and draw a combination of these views, as in the photos on my Figs 216 and 219. Judging by his figures and descriptions, at that time he did not see eggs of P. macronyx, and the female described by him as Centroptilum nana is not conspecific with P. macronyx .</p><p>In another publication (Bogoescu &amp; Tabacaru 1966), Centroptilum nana was included in a key for female imagines, and its egg was characterized as «Die Äquatorialzone des Chorions mit 5–7 Warzenreihen» (the equatorial zone of chorion with 5–7 rows of warts). This wordily characteristics agrees with P. macronyx, but figure was not given, and material examined was not reported.</p></div>	https://treatment.plazi.org/id/039187C0FFF6FFDA83CFF88D37E366AA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Kluge, Nikita J.	Kluge, Nikita J. (2025): Contribution to the knowledge of psammophylous representatives of Procloeon Bengtsson 1915 - Psammonella Glazaczow 1987 and Pseudocentroptiloides Jacob 1987 (Ephemeroptera: Baetidae). Zootaxa 5691 (3): 449-494, DOI: 10.11646/zootaxa.5691.3.3, URL: https://doi.org/10.11646/zootaxa.5691.3.3
