Alainites albai El Alami, Vuataz & Gattolliat sp. nov.

Figs 1, 2, 3, 4, 5, 6, 7, 8

Type-material.

Holotype. Morocco. Nymph; Chefchaouen Province, S10 Oued Kelâa; Loc. Akchour; 35°14'32"N, 05°10'10"W; alt. 460 m; 3.iii.2021; El Yaagoubi leg.; GBIFCH00763782; MZL.

Paratypes. Morocco. 1 nymph; same data as holotype; DNA; GBIFCH00980877; MZL • 2 nymphs; same data as holotype; 7.xi.2014; El Bazi leg.; in alcohol; LESCB • 1 nymph; Chefchaouen Province, S1 Oued Amazithen, Loc. El Ouesteyine; 35°17'57.264"N, 4°54'33.732"W; alt. 166 m; 27.ii.2021; El Yaagoubi leg.; DNA; GBIFCH01144392; MZL • 2 nymphs; Chefchaouen Province, S2 Oued Sidi Yahia Aarab, Loc. Sidi Yahia Aarab; 35°17'10.428"N, 4°53'37.644"W; alt. 114 m; 28.v.2022; El Yaagoubi leg.; one in alcohol; GBIFCH00763784 and other DNA; GBIFCH01144394; MZL; 1 nymph; 18.vi.2014; Khadri leg.; in Alcohol; GBIFCH00763781; MZL; 4 nymphs; 28/v/2022; El Yaagoubi leg.; LESCB • 6 nymphs; Chefchaouen Province, S3 Oued Jenane Niche, Loc. Jenane Niche; 35°16'44.904"N, 4°51'40.788"W; alt. 93 m; 9.ix.2021; El Yaagoubi leg.; LESCB • 11 nymphs; Chefchaouen, Province, S4 Oued Aârkob, Loc. Arherarose; 35°15'59.4"N, 4°50'33.216"W; alt. 128 m; 9.xii.2021; El Yaagoubi leg.; in alcohol; LESCB • 22 nymphs; Chefchaouen Province, S5 Oued Assifane, Loc. Igourain; 35°7'6.7584"N, 4°59'3.9984"W; alt. 1405 m; 9.ix.2021; El Yaagoubi leg; 1 on slide; LESCB • 5 nymphs; Chefchaouen Province, S6 Oued Beni Mhammed, Loc. Beni Mhammed; 35°09'34.0812"N, 5°07'34.0212"W; alt. 1330 m; 29.v.2008; El Alami leg.; in alcohol; GBIFCH00763777; MZL • 2 nymphs; Chefchaouen Province, S7 Oued Afeska, Loc. Afeska; 35°10'11.0388"N, 5°13'6.2988"W; alt. 1293 m; 2.vi.2021; El Yaagoubi leg.; 1 nymph DNA; GBIFCH01144391; 2.vi.2020;2 in alcohol; GBIFCH00763777; GBIFCH00763783; MZL and 4 nymphs; 2.vi.2022; El Yaagoubi leg.; 3 in alcohol and 1 on slide; LESCB • 10 nymphs; Chefchaouen Province, S8 Oued Maggo, Loc. Maggo Nord village; 35°6'48.6"N, 5°11'26.7"W; alt. 905 m; 24.ii.2022; El Yaagoubi leg.; LESCB; 5 nymphs; 3.vi.2016; El Alami, leg.; in alcohol; GBIFCH00763780; MZL • 4 nymphs; Chefchaouen Province, S9 Oued Ouara, Loc. Khizana; 35°02'614"N, 5°14'016"W; alt. 930 m; 23.vi.2022; El Yaagoubi leg.; 2 on slide; LESCB; 2 for DNA; GBIFCH01144393; GBIFCH01144390; MZL • 3 nymphs; Chefchaouen Province, S11 Oued Laou, Loc. Abiyati; 35°17'55.14"N, 5°13'59.99"W; alt. 140 m; 11.iii.2001; El Alami leg.; in alcohol; GBIFCH00763779; MZL • 1 nymph; Chefchaouen Province, S12 Oued Harakat, Loc. Mezine village; 35°6'8"N, 5°20'46"W; alt. 740 m; 31.iii.2021; El Yaagoubi leg.; DNA, GBIFCH00980924; MZL • 20 nymphs; Chefchaouen Province, S13 Oued Mansoura, Loc. Tanaqoub; 35°5'16"N, 5°30'37"W; alt. 124 m; 01.vi.2021; El Yaagoubi leg.; 2 on slide; LESCB • 2 nymphs; Ouezzane Province, S14 Oued Loukkos, Loc. Souk El Had; 35°01'21"N, 5°25'14"W; alt. 140 m; 11.iv.2021; El Yaagoubi leg.; in alcohol; GBIFCH00763776; MZL • 1 nymph; Tetouan Province, S15 Oued Tisgris, Loc. Hammadesh; 35°22'02"N, 5°32'02"W; alt. 505 m; 20.iv.2021; El Yaagoubi leg.; DNA; GBIFCH01144254; MZL • 10 nymphs; Tetouan Province, S16 Oued Rmel, Loc. Ain Dchicha; 35°52'40"N, 5°28'24"W; alt. 49 m; 20.x.1997; El Alami leg.; in alcohol; LESCB and 2 nymphs in alcohol; GBIFCH00763778; MZL.

Differential diagnosis.

Alainites albai sp. nov. can be distinguished from other West Palaearctic species of Alainites based on the combination of nymphal characters, summarized in Table 2: (a) seven pairs of abdominal gills, (b) paraproct prolongation covered with small spines on its entire surface with broad, triangular spines laterally, (c) small teeth between prostheca and mola on both mandibles, (d) low number of dorsal setae on its fore-femora (10-15).

Morphological description.

Nymph. Length. Female body 6.0-7.9 mm; cerci 4.5-5.5 mm; median caudal filament 1.3-1.4 mm (ca 2/3 of cerci); Male body 6.0-6.7 mm; cerci 4.0-5.0 mm; median caudal filament ca 2/3 of cerci.

Coloration.

General coloration pale to medium brown (Figs 2, 3). Head uniformly pale brown to brown with yellow vermiform marks on vertex and frons (Fig. 3a). Turbinate eyes in male nymph’s purple-brown. Legs ecru except upper side of femora brown (Fig. 2b). Thorax with some paler clear pattern (Fig. 3c). Abdominal tergites pale brown with a central, elongated, yellowish dot; distal part of tergite IX and whole tergite X pale brown to yellowish. Abdominal sternites yellowish to pale brown. Cerci ecru to pale brown without bands or pattern.

Head. Antennae (Fig. 3a) close to each other, with a narrow inter-antennal carina, scape with deep scale insertions, pedicel with deep scale insertions and few setae (Fig. 3b).

Dorsal surface of labrum (Fig. 3d) with one central long seta and distolateral arc of two long, simple, stout setae, and small fine setae scattered on surface; ventral surface with 5-8 submarginal small, pointed setae; distal margin fringed with ca 20 short, followed by eight or ten long, feathered setae.

Right mandible (Fig. 4a) shagreened, with scattered fine setae and deep scale insertions; incisors composed of eight blunt, distinct denticles, outer- and inner denticles notably smaller than others; prostheca reduced and bifid with numerous thin setae; 10-16 teeth in the space between prostheca and mola (Fig. 4b), tuft of setae absent.

Left mandible (Fig. 4c) slightly shagreened, with sparse fine setae and deep scale insertions; incisors composed of seven blunt, distinct denticles, outer denticle conspicuously smaller than others; prostheca with one row of 11-15 medium denticles and a comb-like structure (Fig. 4d); edge between prostheca and mola with sparse spines particularly in apical half, without setae.

Hypopharynx (Fig. 5a) trilobed, apically covered with thin setae; lingua with small central protuberance; superlingua longer than lingua.

Maxilla (Fig. 5b) apex with three elongated acute and curved teeth and a toothlike dentisetae; crown with one row of small setae ending with stouter and longer ones (Fig. 5c); palp two-segmented, extending behind apex of galea-lacinia, length of segment I approximately 0.65 × segment II; segment II apically rounded, with few thin setae.

Labium (Fig. 5d) with glossae slightly shorter than paraglossae; inner margins of glossae with 7 or 9 stout medium setae, apical margin with ca 5-7 long stout setae, ventral surface with few thin scattered setae; dorsal surface with row of ca 6-9 medium setae; paraglossae of constant width, with three rows of 11 or 14 long, stout and feathery setae apically; labial palp three-segmented; segment I 0.8 × length of segments II and III combined; segment II with dorsal oblique row of 4-6 medium setae; segment III conical, nearly symmetrical, slightly pointed apically, covered with few medium stout setae and small pointed setae apically.

Thorax. Forelegs (Fig. 6): Trochanter with seven marginal short stout pointed setae; femur dorsal margin with one regular row of 10-15 long, robust setae, and two subapical stout and long setae; ventral margin with short stout pointed setae; lateral surface with sparse scale bases, mainly on apical half and along subdorsal area (Fig. 6a). Tibiae (Fig. 6b) dorsally with 6-10 short stout pointed setae and single apical seta; ventrally with marginal and submarginal short stout pointed setae, denser towards apical end; tibiopatellar suture absent; lateral surface with few short, stout, pointed setae and numerous scale bases. Tarsi bares dorsally; ventral margin with ca 15-24 pointed medium setae; lateral surface with numerous scale bases. Tarsal claws (Fig. 6c) hooked with one row of 11-13 medium teeth, apical setae absent (Fig. 6d). Mid and hindlegs similar to forelegs except femora dorsally with 13-17 pointed setae and tibiae with 21-24 setae dorsally.

Hindwing. Pads present.

Abdomen. Terga (Fig. 7a) shagreened, with numerous scale bases, distal margin of tergite IV with triangular spines ~ 2 × longer than broad. Sternites with scale bases; posterior margin smooth without spination.

Gills (Fig. 2d) on segments I-VII, elliptic, almost symmetrical and serrated all along costal and distal margins except proximal part and with few setae, gill I smaller than others, length of gill I longer than half of tergite II; gill IV (Fig. 7b) and gill V larger than others, (Fig. 7c). Surface with pores and without any setae; tracheation well visible but with reduced ramification. Paraproct (Fig. 7d, e) shagreened with scale bases and setae; margin with seven broad, triangular spines inner to prolongation and numerous medium spines outer to prolongation; lateral margins of prolongation with numerous medium to broad spines, surface covered with spines; cercotractor with scattered scale bases, distal margin with ca 20 medium spines.

Imago. Unknown.

Molecular analysis.

The COI data set was> 95% complete and included 35% of parsimony informative sites. Pairwise COI distances across all sequences ranged from 0% to 23.4%. The overall mean p-distance within MOTUs was 0.8% (mean range: 0.1%-2.3%), while the overall mean p-distance between MOTUs was 18.7% (mean range: 14.9%-22.6%). The maximum p-distance within MOTUs varied from 0.2% ( A. gasithi and A. kars) to 2.8% ( A. sp. 2). The minimum p-distance between MOTUs ranged from 14.3% ( A. sp. 1- A. sp. 2) to 16.7% ( A. kars - A. gasithi and A. kars - A. sp. 4). The seven sequences from A. albai sp. nov. formed a strongly supported monophyletic clade, identified as a distinct MOTU in the three species delimitation analyses (Fig. 8). All methods were fully congruent in delimitating the other MOTUs as well.

Distribution

(Fig. 1). Morocco.

Etymology.

The first author dedicates the name of this species to her former mentor, Professor Javier Alba-Tercedor, a specialist in the Ephemeroptera of the Iberian Peninsula. He contributed significantly to her training and specialization in the taxonomic study of the Ephemeroptera of Morocco.