Carinogalumna erciyesi sp. nov.

(Figs 1–19)

Diagnosis

Adults brown to dark brown, medium-sized (668–750), with characters of Carinogalumna given by Ermilov and Klimov (2017). Interlamellar seta long, bothridial seta setiform, with narrow, barbed head. Notogaster with 10 pairs of alveolar setae and four pairs of porose areas, with largest Aa, most rounded or oval, but A2 and A3 can also be elongated. Porose area A1 located slightly closer to seta h 3 than to lp. Three pairs of genital setae in anterior position, postanal porose area elongated.

Juveniles light brown with darker prodorsal, gastronotal and genital shields and epimeres. In all juveniles prodorsal setae of medium size or long and barbed, and bothridial seta clavate, with long, narrow, barbed head. Larva with 12 pairs of gastronotal setae, including alveolar h 3, nymphs with 15 pairs; most setae of medium size and barbed, except for short p 2 and p 3, smooth in protonymph; setae p 2 and p 3 inserted on unsclerotized integument. Gastronotal shield well-developed, with setae of d -, l -series and h 1 in larva, and of d -, l -, h -series and p 1 in nymphs. In all juveniles, setae of c -series inserted on microsclerites, and genital opening located on large sclerite. In all juveniles, humeral organ present, with porose area above it.

Description of adult

Measurments. Body length (and range) of females 719.9±28.1 (685–750, n= 15), and males 684.6±17.9 (668– 701, n= 15), body width (and range) of females 499.9±8.4 (489–505), and males 481.9±8.9 (473–489).

Cerotegument. Brown, microporose, with dense, short lines on dorsal part of pteromorphs, delicate stria and light, oval spots from pteromorphs to medial part of notogaster and several pits between setal pair h 1.

Prodorsum. Rostrum rounded (Figs 1a, 2, 5a, 5c, 7d) and pointed in lateral aspect (Figs 4a, 4b 5b, 5d). Rostral setae of medium size (ro, 83–87), inserted on lateral part of rostrum, lamellar setae longer (le, 140–147) inserted behind ro and medial to lamellar line (L), both finely barbed (Figs 1a, 4a, 4b). Interlamellar setae long (in, 98–102) and barbed. Mutual distance between setal pair le slightly longer than that between setal pair ro, and mutual distance between setal pair in similar to that between setal pair ro. Bothridium (bo) oval, hidden under anterior edge of notogaster, connected with pteromorph. Bothridial setae long (bs, 141–145), with barbed head (Figs 1a, 1c, 4a, 5a, 5b, 5d, 6a, 6b), exobothridial setae of medium size (ex, 26) and barbed. Porose area Ad elongated (32 x 8), dorsosejugal suture complete.

Notogaster. Almost spherical, with 10 pairs of alveolar setae including c 2 on pteromorphs (Figs 1a, 3, 4a, 5a, 5b, 6b–d). Four pairs of porose areas present; Aa largest, oval (35 x 13), A1 and A2 smaller, usually oval (15 x 10), and A3 usually elongated (30 × 18), but A2 can also be elongated (23 x10) and A3 can be oval as A1 (Figs 1a, 3, 4a, 5a, 5b, 6b–d). Lyrifissure ia on medial part of pteromorph, im anteromedial to seta lp, approximately in similar distance from seta lm and lp, ip lateral to seta p 1, and ih and ips anterior and anterolateral to seta p 3, respectively; opisthonotal gland opening gla anterior to seta h 3 (Figs 1a, 4a).

Ventral side. Subcapitular setae a, m and h short and smooth (12–16, Figs 2, 5c, 7d); chelicera chelate-dentate (201 x 83), cheliceral seta cha longer than chb, both barbed; palp relatively short (125), most palp setae barbed (Figs 4c, 4d, 7d), formula of palp setae (femur to tarsus and solenidion): 0-2-1-3-9(1). Epimeral setae short (6) and thin, formula of epimeral setae 1-1-2-1. Genital setae longer (6 pairs, 18–25) than epimeral setae, anterior edge of genital plate with three setae (Figs 2, 5c, 7a). Aggenital setae short (10), located closer to genital opening than to anal aperture, adanal setae longer (16–20), all smooth (Figs 2, 5c, 7c). Discidium triangular, circumpedal carina short. Adanal lyrifissure located diagonal and close to anal plate, postanal porose area elongated (120 x 10, Figs 2, 3).

Legs. All leg setae barbed (Figs 5, 6a, 8). Formulae of leg setae (and solenidia), trochanter to tarsus: I—1-4- 3(1)-4(2)-20(2), II—1-4-3(1)-4(2)-15(2), III—1-2-1(1)-3(1)-15; IV—1-2-2-3(1)-12. Leg tarsi heterotridactylous. On prodorsum, notogaster and legs no distinct differences present between females and males.

Description of juveniles

Larva egg-shaped in dorsal and ventral view (Figs 9a, 10a) and unpigmented, except for light brown prodorsal and gastronotal shields, epimeres and legs. Prodorsum subtriangular, prodorsal seta in long, other setae of medium size, ro longest, ex shortest (Figs 9a, 11a, 12, 13a, 13b, Table 1), all barbed. Mutual distance between setal pair le similar as that between setal pair ro, and mutual distance between setal pair in about 1.5 times longer than that between pair ro, pair le inserted clearly closer to pair in than to ro. Opening of bothridium rounded, bothridial seta clavate, with narrow, barbed head (Figs 9a, 11a, 12, 13a–c). Ridges present from bothridium in direction of seta in and ex.

Gastronotum of larva with 12 pairs of setae, including alveolar h 3 located lateral to medial part of anal valves; most medium-sized and barbed except for short h 2 with small barbs (Figs 9a, 10a, 11a, 12a, 12b, 13a, 13b, 13d, 14a). Setae of c -series inserted on microsclerites, length slightly increasing from c 1 to c 3 (Table 1). Gastronotal shield with seven pairs of setae (d -, l -series, h 1), seta h 2 and h 3 inserted on unsclerotized integument. Small porose areas present, Aa anteromedial to seta la, A1 anterior to seta lm, and A2 posteromedial to seta lp (Figs 9a, 11a). Cupule ia located posterolateral to seta c 3, im anterior to seta lp, ip posteromedial to seta h 2, and ih lateral to anterior part of anal valves. Opisthonotal gland opening gla lateral to seta lp, without dark sclerotized surrounding (Figs 10a, 11a). Humeral organ rounded, located above insertions of leg II, and porose area present above this organ. Paraproctal valves (segment PS) glabrous. Legs of larva stocky, all femora flattened, with ventral keels, most leg setae barbed (Figs 12, 13, 14c, 14d, 15a, 16).

Shape of prodorsum of protonymph and other nymphs, prodorsal setae, bothridium and bothridial seta as in larva, but head of bothridial seta slimmer than in larva (Figs 11b, 15b, 15c). Gastronotum of nymphs oval with 15 pairs of setae because setae of p -series appearing in protonymph and retained in subsequent nymphs; most of medium size and barbed except for short p 2 and p 3, smooth in protonymph (Figs 10b, 11b, 15b–d, 17, 18, Table 1). Setae of c -series inserted on microsclerites, length slightly increasing from c 1 to c 3 (Figs 11b, 18). Gastronotal shield with 10 pairs of setae (d -, l -, h -series, p 1), setae p 2 and p 3 inserted on unsclerotized integument (Figs 10b, 11b, 17). Four pairs of porose areas present, Aa anteromedial to seta la, A1 posteromedial to seta lm, A2 medial to seta h 3 and A3 posteromedial to seta h 2 (Figs 11b, 18). In protonymph one pair of genital setae appearing lateral to medial part of genital valves, two pairs added, each in deutonymph and tritonymph (Figs 10b, 17), all short and smooth. In deutonymph, one pair of aggenital setae and three pairs of adanal setae appearing, and retained in subsequent instars; all short and smooth, setae of ad -series inserted on unsclerotized integument. In protonymph and deutonymph, anal valves glabrous, in tritonymph with two pairs of short and smooth anal setae (Figs 10b, 17). In nymphs cupules ia and im as in larva, cupule ip lateral to seta p 1. Only cupules located lateral to anterior part of anal valves visible in punctated integument, ips in protonymph and iad in deutonymph and tritonymph (Figs 10b, 17). Opisthonotal gland opening gla anterolateral to seta h 3, without dark sclerotized surrounding. Humeral organ oval, located as in larva, with porose area above this organ (Fig. 11b). Legs of tritonymph stocky, all femora flattened with ventral keels, all leg setae barbed (Figs 15b–d, 19).

Summary of ontogenetic transformations

In all instars the prodorsal seta le is longest and ex is shortest, but in the larva ro is slightly longer than in, whereas in other instars in is longer than ro. The bothridium is rounded in all instars, but in the adult it is covered by anterior edge of notogaster. In all juveniles the bothridial seta is clavate, with narrow, barbed head, but in the adult, it is setiform, with barbed head. The larva has 12 pairs of gastronotal setae, including alveolar h 3, the nymphs have 15 pairs (p -series appears in the protonymph and is present in other instars), whereas the notogaster of the adult loses all setae and alveoli of setae c 1, c 3 and d -series, such that 10 pairs of alveoli remain. The formula of gastronotal setae in C. erciyesi is 12-15-15-15-10 (from larva to adult, including alveolar setae), the formulae of epimeral setae are 3-1-2 (larva, including scaliform 1c), 3-1-2-1 (protonymph), 3-1-2-2 (deutonymph and tritonymph), and 1-1-2-1 (adult). The formula of genital setae is 1-3-5-6 (protonymph to adult), and formula of aggenital setae is 1-1-1 (deutonymph to adult). Setal formula of segments PS–AN is 03333-0333-022. The ontogeny of leg setae and solenidia of C. erciyesi is as in Pilogalumna tenuiclava given by Seniczak and Seniczak (2022), except for the seta v 1 on genua I and II which in C. erciyesi is present in the tritonymph and adult, and in Pil. tenuiclava it is absent.

Ecology and biology

Carinogalumna erciyesi was found in the pine litter under pine tree in the campus of Erciyes University in Kayseri (Turkey). In this sample C. erciyesi achieved the density of 152 individuals per 500 cm 3, the juveniles constituted 86% of the population of this species. The stage structure of C. erciyesi in two samples was the following: 107 larvae, 105 protonymphs, 40 deutonymphs, 10 tritonymphs and 41 adults. Based on 30 randomly selected adults, the sex ratio (females to males) was 1:1, and 83% of females were gravid and carried 1–4 large eggs (281 x 162 each), comprising 39% of the length of females.

Type material

The holotype (female) and five paratypes (three females and two males) with the above collection data are deposited in the University Museum of Bergen, University of Bergen, Bergen, Norway (ZMUB).

Etymology

The species name erciyesi refers to the Mt. Erciyes, which is located relatively close to the town Kayseri in Turkey, where this species was found.

Comparison of morphology of Carinogalumna erciyesi sp. nov. with congeners and remarks

In Table 2, we compared selected morphological characters of adults of C. erciyesi sp. nov. with those of other Carinogalumna species listed by Subías (2004, 2022). Among these species the largest is C. clericata (Berlese, 1914) and smallest one is C. alineata Ermilov & Martens, 2014. In the light of this comparison, C. erciyesi is most similar to C. montana Engelbrecht, 1973, but differs from it mainly by the shape of the bothridial seta and porose areas Aa and A2. In C. erciyesi, the bothridial seta is setiform (versus clavate in C. montana), and porose areas Aa and A2 are oval, whereas in C. montana they are elongated.

The nymphs of C. erciyesi have most gastronotal setae of medium size, which is unique in known nymphs of Galumnidae. In Table 3, we compared some morphological characters of juveniles of C. erciyesi with those of Galumna flagellata Willmann, 1925, Pilogalumna kazakhstanica Seniczak et al., 2023, Pil. tenuiclava (Berlese, 1908) and Psammogalumna iranica Akrami et al., 2011. These juveniles differ from one another mainly in the presence of humeral organ, microsclerites and the shape of some prodorsal and gastronotal setae. The juveniles of C. erciyesi are most similar to those of G. flagellata, differing from them in six morphological characters, and they are least similar to Pil. tenuiclava, differing from them in 11 morphological characters. The most important character that differentiates C. erciyesi from other species of Galumnidae is the shape of most gastronotal setae of the nymphs. In most species these setae are short (Sengbusch 1954; Woodring 1965; Seniczak & Seniczak 2007, 2022, 2023; Seniczak et al. 2012, 2021, 2022; Bayartogtokh & Ermilov 2017), but in C. erciyesi they are of medium size. Generally, the juveniles of C. erciyesi are similar to those of other species of Galumnidae. Among the above mentioned species, the juveniles of C. erciyesi have a clear porose area located above a humeral organ, but in species having a humeral organ this porose area is absent. A similar porose area is present in G. flagellata, but in this species a humeral organ is absent.

The femora of juveniles of C. erciyesi and G. flagellata are flattened, with ventral keels, whereas those of other species are oval in cross section, without ventral keel. The flattened femora with ventral keels are common in the juveniles of Melanozetes Hull, 1916 (Behan-Pelletier 1985, 1986, 2000; Seniczak et al. 1990, 2015; Seniczak & Seniczak 2018) except for M. avachai Seniczak et al., 2016. This suggests that the shape of femora of juveniles of Galumnidae and Melanozetes is species-specific.