Unravelling the mishmash: A new phylogeny for the family Empheriidae (Psocodea, Trogiomorpha) with a new genus and species from Cretaceous Charentese amber Author Alvarez-Parra, Sergio https://orcid.org/0000-0002-0232-1647 State Key Laboratory of Palaeobiology and Petroleum Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China & Departament de Dinamica de la Terra i de l'Ocea, Facultat de Ciencies de la Terra, Universitat de Barcelona, c / Marti i Franques s / n, 08028, Barcelona, Spain sergio.alvarez-parra@ub.edu Author Nel, Andre https://orcid.org/0000-0002-4241-7651 Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain Author Perrichot, Vincent https://orcid.org/0000-0002-7973-0430 Institut de Systematique, Evolution, Biodiversite (ISYEB), Museum national d'Histoire naturelle, CNRS, Sorbonne Universite, EPHE, Universite des Antilles, CP 50, 45 rue Buffon 75005 Paris, France Author Jouault, Corentin https://orcid.org/0000-0002-3680-5172 Institut de Recerca de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain & Institut de Systematique, Evolution, Biodiversite (ISYEB), Museum national d'Histoire naturelle, CNRS, Sorbonne Universite, EPHE, Universite des Antilles, CP 50, 45 rue Buffon 75005 Paris, France & Univ Rennes, CNRS, Geosciences Rennes, UMR 6118, F- 35000 Rennes, France text Arthropod Systematics & amp; Phylogeny 2024 2024-03-22 82 183 199 http://dx.doi.org/10.3897/asp.82.e114849 journal article http://dx.doi.org/10.3897/asp.82.e114849 1864-8312-82-183 E45565A395804B06AB8B093CB0692937 05355FC72CC05232AC03F5909266BD42 Santonipsocus mimeticus sp. nov. Figures 1 , 2 , 3 , 4 Holotype. MNHN.F.A30180 (ARC-186.7), adult specimen, probably male (Figs 1 - 3 ), erroneously figured as a paratype of Proprionoglaris guyoti in the PhD dissertation of Perrichot (2005 , fig. 31b). Amber fragment cut from a larger amber piece ( 'ARC-186' ) and prepared in Canada balsam. Originally in syninclusion with the holotype of Prospeleketor albianensis Perrichot, Azar, Neraudeau and Nel, 2003 (specimen MNHN.F.A30111 [ARC-186.10]) and nine other arthropods ( Diptera , Hemiptera , Coleoptera , and Pseudoscorpionida ). In the article by Perrichot et al. (2003) , it is indicated that the holotype of P. albianensis is together with the female paratype of P. guyoti as syninclusions. However, P. guyoti was described from the female holotype (MNHN.F.A30108 [ARC-58.2]) and two male paratypes (MNHN.F.A30109 [ARC-50.1] and MNHN.F.A30110 [ARC-201.2]), all in other amber pieces. All the above-mentioned material is housed in the palaeontological collection of the Museum National d'Histoire Naturelle, Paris, France. Figure 1. Santonipsocus mimeticus gen. et sp. nov. ( Psocodea , Trogiomorpha , Empheriidae ), holotype MNHN.F.A30180 (ARC-186.7), probably male; uppermost Albian-lowermost Cenomanian amber of Archingeay-Les Nouillers (France). A habitus from dorsal view; B habitus from ventral view. Both at same scale. Figure 2. Santonipsocus mimeticus gen. et sp. nov. ( Psocodea , Trogiomorpha , Empheriidae ), holotype MNHN.F.A30180 (ARC-186.7), probably male; uppermost Albian-lowermost Cenomanian amber of Archingeay-Les Nouillers (France). A photograph of right fore- and hind wings, overlapped; B schematic drawing of forewing; C schematic drawing of hind wing. B and C at same scale. Figure 3. Santonipsocus mimeticus gen. et sp. nov. ( Psocodea , Trogiomorpha , Empheriidae ), holotype MNHN.F.A30180 (ARC-186.7), adult specimen, probably male; uppermost Albian-lowermost Cenomanian amber of Archingeay-Les Nouillers (France). A photograph of left antenna; B detail of flagellomeres of left antenna; C head from ventral view; D , E photograph and schematic drawing, respectively, of nodulus area of right forewing, both at same scale, blue arrowheads: meeting point of veins Cu2 and 1A with forewing margin, red arrowhead: emergence of R1 in relation to basal cell in hind wing; F photograph of left midleg and hind leg; G photograph of tarsus of right hind leg; H genitalia in ventral view. Other material. IGR.ARC-169, adult specimen, sex unknown (Fig. 4 ). From Archingeay-Les Nouillers amber deposit. Amber piece prepared in Canada balsam; the forewings are separated from the rest of the body. Undetermined arthropod leg as syninclusion. Housed in the Geological Department and Museum of the University of Rennes, France . Figure 4. Santonipsocus mimeticus gen. et sp. nov. ( Psocodea , Trogiomorpha , Empheriidae ), specimen IGR.ARC-169, sex unknown; uppermost Albian-lowermost Cenomanian amber of Archingeay-Les Nouillers ( France ). A habitus from dorsal view; B habitus from ventral view; C photograph of wings; D schematic drawing of forewing; E photograph of the tarsi of right foreleg and midleg. Locality and horizon. Font-de-Benon quarry, Archingeay-Les Nouillers, Charente-Maritime Department (Nouvelle-Aquitaine, France); level A1sl, uppermost Albian-lowermost Cenomanian, Cretaceous ( Neraudeau et al. 2002 ). Etymology. From the Greek μιμητικός , meaning 'imitator' , based on the resemblance with other Cretaceous barklice species. Diagnosis. As for the genus (vide supra). Description. Probably male. Body length 1.75 mm from clypeus to genitalia (Fig. 1 ). - Head : almost twice wider than long, 0.26 mm long, 0.44 mm wide; vertex covered by setae; marked epicranial suture with indistinct anterior arms; no ocelli; compound eye not prominent and small; left antenna complete with 17 flagellomeres (Fig. 3A ), right antenna with eight flagellomeres preserved; short setae on proximal flagellomeres; scape 0.04 mm long, pedicel 0.05 mm long; lengths of flagellomeres of left antenna: f1 0.04 mm, f2 0.04 mm, f3 0.05 mm, f4 0.07 mm, f5 0.07 mm, f6 0.06 mm, f7 0.06 mm, f8 0.07 mm, f9 0.06 mm, f10 0.07 mm, f11 0.07 mm, f12 0.04 mm, f13 0.06 mm, f14 0.06 mm, f15 0.05 mm, f16 0.04 mm, f17 0.04 mm; flagellomeres lacking secondary annulations (Fig. 3B ); clypeus bulging; maxillary palps four-segmented, covered by short setae (Fig. 3C ); lengths of maxillary palpomeres: I 0.02 mm, II 0.08 mm, III 0.02 mm, IV 0.07 mm; sensillum not present on second maxillary palpomere; distal maxillary palpomere globose and rounded; labial palps seemingly two-segmented, with the distal labial palpomere showing an elongate shape 0.06 mm long (Fig, 3C); a structure might correspond to lacinia. - Thorax : 0.46 mm long; pronotum bulging, covered by a few setae; macropterous (Fig. 2 ). - Wings : membrane hyaline with setae; forewing and hind wing with nearly the same size, surpassing the distal part of the abdomen only slightly (Fig. 1 ); forewing with margin covered by setae, veins with two rows of setae (Fig. 2A ); hind wing glabrous (Fig. 2B ). Forewing 3 x longer than wide, 1.39 mm long and 0.46 mm wide (Fig. 2A ); basal section of Sc curved and joining R1 at 0.60 mm from wing base, forming a narrow cell; a short, curved crossvein emerging from basal section of Sc, very close to the meeting point of Sc and R1, reaching margin at 0.67 mm from wing base; distal bent of basal section of Sc between emerging of crossvein and meeting point with R1 curved, not straight or perpendicular to R1; distal section of Sc separating from R1 and joining wing margin at 1.02 mm from wing base showing a sigmoidal path; R1 long reaching margin at 1.22 mm from wing base; basal section of Rs oblique, with crossvein between R1 and Rs present, forming a six-angled radial cell; vein Rs+M 0.06 mm long; Rs bifurcating at 0.86 mm from wing base; R2+3 and R4+5 reaching margin at 1.34 mm and 1.37 mm from wing base, respectively; bifurcation of M1 and M2 at 0.90 mm from wing base, reaching margin at 1.38 mm and 1.34 mm from wing base, respectively; M3 emerging from M at 0.65 mm from wing base and reaching margin at 1.26 mm from wing base; cells between R2+3 and R4+5, and between M1 and M2, elongate and relatively narrow; bifurcation of Cu1 into Cu1a and Cu1b at 0.48 mm from wing base; elongate and narrow areola postica, with Cu1a extending towards apex; Cu1a and Cu1b reaching margin at 1.11 mm and 0.79 mm from wing base, respectively; evanescent Cu2, without rows of setae, extending straight and reaching margin at 0.65 mm from wing base; vein 1A showing sigmoidal path and reaching margin at 0.56 mm from wing base; no nodulus (Fig. 3D, E ); joining of Cu2 with margin separated by 0.11 mm from joining of 1A with margin. Hind wing almost 3 x longer than wide, 1.20 mm long and 0.42 mm wide (Fig. 2B ); Sc not visible; basal cell closed, elongate and narrow, with three angles, 0.15 mm long and 0.03 mm wide; R1 not emerging from apex of basal cell; R1 fused to Rs+M for 0.03 mm; R1 emerging from basal cell in specimen IGR.ARC-169; sigmoidal R1 reaching margin at 0.83 mm from wing base; bifurcation of Rs into R2+3 and R4+5 at the same level as R1 reaching margin; R2+3 and R4+5 reaching margin at 1.14 mm and 1.19 mm from wing base, respectively; cell between M1 and M2 elongate and narrow; M1 and M2 reaching margin at 1.12 mm and 0.97 mm from wing base, respectively; Cu1 reaching margin at 0.62 mm from wing base; a vein reaching margin close to Cu1 might correspond to Cu2; anal vein not visible. - Legs : femora thick, without setae; tibiae thin, with three to four spines, and covered by setae; tarsi three-segmented, with first tarsomere covered by a few short setae (Fig. 3F, G ); lengths of tarsomeres in foreleg: I 0.10 mm, II 0.04 mm, III 0.03 mm; lengths of tarsomeres in midleg: I 0.13 mm, II 0.03 mm, III 0.03 mm; lengths of tarsomeres in hind leg: I 0.18 mm, II 0.05 mm, III 0.04 mm; pretarsal claws lacking preapical tooth and pulvillus. - Abdomen : 1.03 mm long; genitalia poorly visible, although male characters, such as hypandrium and paraprocts, are discernible (Fig. 3H ). Remarks. Although the specimen IGR.ARC-169 is poorly preserved (Fig. 4A, B ), it is preliminary considered conspecific with holotype MNHN.F.A30180 (ARC-186.7) based on coincident diagnostic characters of the new genus: body length from clypeus to genitalia (1.75 mm holotype vs. 1.72 mm); distal maxillary palpomere globose and rounded; forewing with setae on margin, two rows of setae on veins, and very similar venation (Fig. 4C, D ), particularly presence of a cross-vein between Sc and margin emerging very close to meeting point of Sc with R1, distal bent of basal section of Sc curved between emerging of crossvein and meeting point with R1, not straight or perpendicular to R1, without nodulus; and pretarsal claws lacking preapical tooth and pulvillus (Fig. 4E ). The forewing venation of IGR.ARC-169 is nebulous and difficult to see, but it can be discerned using a combination of transmitted and reflected light under a stereomicroscope. We consider the distal maxillary palpomere globose and rounded and the lack of nodulus as key characters of the new genus. Two differences in wing venation can be related to intraspecific variability or teratism typical in barklice ( Smithers 1972 ): forewing with basal section of Rs placed more proximal in IGR.ARC-169 than in holotype and hind wing with four-angled basal cell in IGR.ARC-169 in comparison with three-angled basal cell in the holotype. Based on the coincident characters and the poor preservation of IGR.ARC-169, we believe that the most parsimonious option is to place this specimen, at least preliminarily, in the same species as the holotype, yet refraining from including it within the type material. Notes on Proprionoglaris guyoti and Proprionoglaris axioperierga . The presence of a nodulus in forewing is confirmed for both species (Fig. 5 ). The species P. guyoti was described as possessing only one row of setae along veins in forewing ( Perrichot et al. 2003 ). However, a detailed re-examination has allowed us to distinguish two rows of setae along veins in all the type specimens. Therefore, the diagnosis must be emended replacing "setae arranged in one row on veins" with "setae arranged in two rows on veins" in forewing. The species P. axioperierga was originally described with two rows of setae along veins in forewing ( Azar et al. 2014 ). Figure 5. Forewings of Proprionoglaris guyoti and Proprionoglaris axioperierga ( Psocodea , Trogiomorpha , Empheriidae ). A holotype of P. guyoti , female, MNHN.F.A30108 (ARC 58.2 R); B paratype of P. guyoti , male, MNHN.F.A30109 (ARC 201.2); C - E new specimens of P. guyoti , sex unknown, IGR.ARC-352.1, IGR.ARC-157, and IGR.ARC-355, respectively; F holotype of P. axioperierga , female, IGR.GAR-69; G paratype of P. axioperierga , sex unknown, IGR.GAR-39. White arrowheads: nodulus.