Two new species of Diamesinae (Diptera: Chironomidae) from the Central Iranian Range and Elburz Mountains (Iran), with new faunistic records for the subfamily Author Ghaderi, Edris Department of Fisheries and Aquatic Ecology, Faculty of Fisheries and Environmental Sciences, Gorgan University of Agricultural Sciences and Natural Resources, Gorgan, Iran & Department of Fisheries Sciences, Faculty of Natural Resources, University of Kurdistan, Sanandaj, Iran Author Namayandeh, Armin Department of Environmental and Life Sciences, Trent University, 1600 West Bank Drive, Peterborough, Ontario, Canada Author Ibrahimi, Halil Department of Biology, Faculty of Mathematics and Natural Sciences, University of Prishtina, “ Hasan Prishtina ”, Prishtina, Kosovo Author Mohammadi, Habibollah Department of Fisheries Sciences, Faculty of Natural Resources, University of Kurdistan, Sanandaj, Iran & Zrebar Lake Environmental Research, Kurdistan Studies Institute, University of Kurdistan, Sanandaj, Iran Author Karimian, Erfan Department of Fisheries Sciences, Faculty of Natural Resources, University of Kurdistan, Sanandaj, Iran & Zrebar Lake Environmental Research, Kurdistan Studies Institute, University of Kurdistan, Sanandaj, Iran Author Mansouri, Arman Department of Fisheries Sciences, Faculty of Natural Resources, University of Kurdistan, Sanandaj, Iran Author Molodi, Farshad Department of Fisheries Sciences, Faculty of Natural Resources, University of Kurdistan, Sanandaj, Iran text Zootaxa 2024 2024-11-07 5537 1 76 94 http://dx.doi.org/10.11646/zootaxa.5537.1.4 journal article 10.11646/zootaxa.5537.1.4 1175-5326 14239318 E45E8227-5CE3-4C26-9E2E-6B65495D9566 Sympotthastia golalae Ghaderi, Namayandeh et Karimian , sp. nov. urn:lsid:zoobank.org:pub: E45E8227-5CE3-4C26-9E2E-6B65495D9566 Figs. 5–7 Type material. Holotype : male; Iran , Kurdistan Province , Central Iranian Range , Bayanchub ; 35.5734466˚N, 46.971835˚E; elev. 1936 m ; 27.iii.2023 ; leg. E. Ghaderi & E. Karimian ; dep. ROM . Paratypes : 3 males same as holotype. 1 male and 1 female ; Iran , Kurdistan Province , Central Iranian Range , Saqqez River ; 36.236831˚N, 46.105522˚E; elev. 1562 m ; 14.iii.2023 ; leg. E. Ghaderi ; dep. ROM . Diagnostic characters. S. golalae sp. nov. can be separated from related species based on the following combination of characters: The adults are very large, with extended wings. Male with AR 3.7–4.0; HW/ PL 0.87– 0.94; sigmoid superior volsella with apically rounded distal part; spiniferous median volsella long and very wide, with apex reaching beyond the 2/3 rd of the superior volsella. Female with HW/ PL 0.88; seminal capsule ovate, with a reduced neck. Etymology. The new species is named in honor of Golale Eshghi, who has tirelessly supported our ongoing studies and research for many years. FIGURE 5. Sympotthastia golalae sp. nov. adult male (A–E). A. antenna. B. head. C. thorax. D. hypopygium. E. phallapodeme, aedeagal lobe, superior volsella, and median volsella. Scale bars are 100 µm. Description Adult male (n = 5). Total length 6.5–7.2, 6.9 mm . Wing 4.3–4.7, 4.6 mm long and 1.1–1.3, 1.2 mm wide. Coloration . Head, thorax, halters and abdomen dark brown, legs slightly lighter. Wing bicolored, with cells m, r, r 1 , and r 4+5 brown and remaining cells light greyish brown. Head ( Fig. 5A–B ). Antenna with 13 flagellomere; last flagellomere with 15–22, 17 sensilla chaetica ( Fig. 1a ), 2 nd –3 rd segments each with 2 sensilla chaetica, groove starts at 4 th segment; subapical setae 32–61, 45 μm long; AR 3.7–4.0, 3.8 ( Fig. 5A ). Eyes bare, with short dorsomedial extension. Temporal setae 8 outer verticals ( Fig. 5B ). Tentorium 272–319, 303 μm long. Clypeus ovate-shaped, without long setae, 85–115, 100 μm long and 133–183, 161 μm wide. Palpal segments lengths (in μm): 89–121, 106; 90–108, 96; 180–194, 186; 227–294, 265; 241–253, 245; second and third palpomere partially fused; third palpomere with 3 sensilla chaetica and circular sensilla capitata at the base, apex with large semicircular sclerotized projection. HW/PL 0.87–0.94, 0.91. FIGURE 6. Sympotthastia golalae sp. nov. adult male. A. wing. B. hypopygium, aedeagal lobe, superior volsella, median volsella and phallapodeme indicated by arrows. C. superior volsella and spiniferous median volsella. Scale bars are 100 µm. Thorax ( Fig. 5D ). Dorsocentrals 26–40, 35 in two to three rows; prealars 15–22, 19; scutellars 50–74, 59 in several rows. Antepronotal lobes with small gap, bearing 7–13, 11 setae. Wing ( Fig. 6A ). Brachiolum with 2 setae. Squama with 83–104, 93 setae. R with 8–16, 13 setae, R 1 with 12–22, 17 setae, other veins bare. Alula bare. Costa extension 140–181, 167 μm long. Anal lobe well-produced. VR = 0.7–0.9, 0.8. Microtrichia only visible at> 100 × magnification. Legs . All legs with long beards. Fore tibia spur 102–138, 115 μm long, mid tibia spurs 86–106, 94 and 69–92, 83 μm long, hind tibia spurs 100–141, 117 and 62–82, 74 μm long, hind tibia comb with around 12–15, 14 spines. Pseudospurs on tarsal segments 1–2 of the mid and hind legs. Lengths and proportions of legs as in Table 4 . TABLE 4. Lengths (in μm) and proportions of leg segments of Sympotthastia golalae sp. nov. , male.

fe	ti	ta 1	ta 2	ta 3	ta 4	ta 5	LR	BV	SV
P 1	1649	1821	1223	635	441	249	217	0.7	3.1	2.9
P 2	1758	1812	803	480	335	205	206	0.4	4.4	5.0
P 3	1765	1910	1193	632	442	242	174	0.6	3.3	3.1

Hypopygium ( Figs. 5D , 6B–C ). Tergite IX with 18 long setae on each side. Laterosternite IX with 11–12 long setae on each side. Anal point absent. Sternapodeme 217–256, 243 μm long, M-shaped, thick, and without oral projections. Phallapodeme 125–230, 195μm long. Sigmoid superior volsella 126–281, 180 μm long, distal part apically rounded. Spiniferous median volsella 102–163, 127 μm long; very wide, with apex reaching beyond the 2/3 rd of the superior volsella. Gonocoxite 413–553, 480 μm long. Inferior volsella a well-reduced lobe. Gonostylus 253–305, 280 μm long, gradually expanded anteriorly; megaseta 26–37, 32 μm long. HR 1.4–2.0, 1.7; HV 2.4–2.8, 2.5. Adult female (n = 1). Total length 7.1 mm . Wing 5.1 mm long and 1.7 mm wide. Color . Same as male. Head ( Fig. 7A ). Antennal flagellomeres 1–2 (in μm): 167, 78; remaining flagellomeres missing. Eyes bare, with short dorsomedial extension. Temporal setae 12, including 6 orbitals and 6 frontals. Tentorium 313 μm long. Clypeus ovate-shaped, without setae, 164 μm long and 234 μm wide. Palpal segments lengths (in μm): 135, 111, 183, 287, 290; second and third palpomere partially fused; third palpomere with 3 sensilla chaetica and circular sensilla capitata at the base, apex of third palpomere with large semicircular sclerotized projection. HW/ PL 0.88. Thorax ( Fig. 7B ). Dorsocentrals 57 in two to three rows; prealars 30; scutellars 100 in several rows.Antepronotal lobes with small gap, bearing 12 setae. Wing ( Fig. 7C ). Brachiolum with 2 setae. Squama with 91 setae. R with 9 setae, R 1 with 15 setae, other veins bare. Alula with 34 setae. Costa extension 226 μm long. Anal lobe well-produced. VR = 0.7. Microtrichia only visible at> 100 × magnification. Legs . All legs with long beards. Fore tarsus missing; fore tibia spur 112 μm long, mid tibia spurs 112 and 107 μm long, hind tibia spurs 142 and 111 μm long, hind tibia comb with around 15 spines. Pseudospurs on tarsal segments 1–2 of the mid and hind legs. Lengths and proportions of legs as in Table 5 . TABLE 5. Lengths (in μm) and proportions of leg segments of Sympotthastia golalae sp. nov. , female.

fe	ti	ta 1	ta 2	ta 3	ta 4	ta 5	LR	BV	SV
P 1	1653	1851	-	-	-	-	-	-	-	-
P 2	2356	1400	691	449	248	263	263	0.6	3.2	3.1
P 3	2175	2356	1400	691	449	248	263	0.6	3.6	3.2

Genitalia ( Figure 7D–E ). Gonapophysis VIII divided into large, semicircular ventrolateral lobe covering the dorsomesal lobe; apodeme lobe distinct, without tubercle, slightly covered by ventrolateral lobe ( Fig. 7D ). Labium inner section well-sclerotized. Seminal capsule 183, 188, 191 µm long and 170, 193, 164 µm wide, respectively; ovate, with a reduced neck; spermathecal duct not visible in this specimen. Notum 172 µm long, with ramus 289 µm long. Tergite IX large, undivided, apex rounded, setigorous protrusion bearing 26 setae. Gonocoxite slightly extended bearing 13 setae. Cercus hexagonal, 360 µm long and 336 µm wide ( Fig. 7E ). Remarks. S. golalae sp. nov. , resembles and most likely is related to Sympotthastia zavreli Pagast 1947 . However, the combination of a higher antennal ratio, the complete absence of an anal point, and larger median volsella separates the adult males of the two species. Sympotthastia wuyiensis Liu, Ferrington et Wang, 2016 is another species without an anal point. However, S. golalae sp. nov. , differs in the shape of superior volsella and the presence of megaseta compared to S. wuyiensis . The prominent adult characteristics of the S. golalae sp. nov. that separate it from other known Sympotthastia species is a larger size and a longer wingspan, which, on average, is 1.3 × for total length and 1.2 × for wing length. Further, a much larger median volsella of S. golalae sp. nov. in comparison to other Sympotthastia species is apparent. The adult female of S. golalae sp. nov. is easily distinguishable from the only species described as female, Sympotthastia bicolor ( Tokunaga, 1937 ) , by a much larger size, and short and wide ovate seminal capsules with small necks. S. bicolor has elongated ovate seminal capsules with prominent necks ( Tokunaga 1937 : Fig. 97). The adult male of Sympotthastia can also be separated based on the key we provided in this study. In constructing this key, we used the keys constructed by Doughman 1985 , Liu et al. (2016) , and Makarchenko (1985) with additions and modifications. In addition to this key, we have listed some relevant distinguishing characteristics of Sympotthastia adult males in Table 6 . FIGURE 7. Sympotthastia golalae sp. nov. adult female (A–E). A. head. B. thorax. C. wing. D. genitalia, ventral. E. genitalia, dorsal. Scale bars are 100 µm. TABLE 6. Comparison of some relevant characteristics of Sympotthastia Pagast, 1947 species adult male. * See notes in the remarks section.

S. diastena	S. fulva	S. gemmaformes	S. golalae sp. nov.	S. huldeni	S. macrocera *	S. repetina	S. spinifera	S. takatensis	S. wuyiensis	S. zavreli
TL (mm)	5.7–6.0	4.7–4.9	3.3	6.5–7.2	5.5	5.5	6–6.5	4.3–4.8	4.5–6.8	4.0–4.3	4–6
WL (mm)	3.6	3.3–3.4	2.8	4.3–4.7	3.6	4.3	3.8–4.0	3.4–3.6	3.9–5.3	2.6–2.8	3–4
HW/PL	-	1.0–1.1	0.93	0.87–0.94	-	-	0.85–0.94	0.79–0.85	0.72–0.90	0.86	1.0–1.1
AR	2.7–2.9	2.3–2.6	3.0–3.4	3.7–4.0	2.8	3.0	2.3–3.1	1.6–1.8	2.8–4.3	1.8	2.3–2.7
TS	-	28–37	19–26	8	10–11	-	31–45	13	15–26	8–18	40–50
Ant	14	15–20	9–12	7–13	-	-	15–17	-	5–15	7–11	15–20
DC	25–30	35–43	25–28	26–40	10–12	-	23–33	22–27	17–24	13–16	35–43
Pa	23	19–26	23–25	15–22	22	-	23–31	12–15	11–24	8–9	8–10
Sq	-	28–31	-	83–104	50–60	-	74–75	-	60–82	44	-
VR	0.88	0.86–0.96	-	0.7–0.9	-	-	-	0.87–0.90	~0.83–0.9	0.83–0.87	0.83
CE L	-	-	-	140–181	-	-	-	-	100–150	113–125	-
LR 1	0.88	0.76	0.84	-	0.84	-	0.82	0.75	0.68–0.82	0.82	0.75
LR 2	0.54	0.48	0.54	0.64	0.56	-	0.48	0.52	0.48–0.51	0.54	0.48
LR 3	0.52	0.57	0.63	0.59	0.70	-	0.6	0.64	0.61–0.64	0.57	0.56
AnP	Naked, awl–like	Weak to strong/ Spiniform	Small tubercle	Absent	Small, narrow & spiniform	Spiniform with 2 bristles	Weak to strong/ Spiniform	Spiniform	Spiniform	Absent	If present, a small tubercle
Anp L	70–90	21–65	-	-	52	-	-	30–40	28–60	-	21–65
Ph L	70–170	-	155	125–230	-	-	-	-	172–215	160–163	-
AL + SVo L	~217	-	221	126–163	-	-	-	-	160–228	-	-
AL + Svo shape	Sigmoid	Apex straight	Reniform	Sigmoid	Sigmoid with apex golf club-shaped	Sigmoid	Reniform	Sigmoid	Sigmoid with apex golf club-shaped	Straight with apex cane– shaped	Apex straight
Ms L	-	29	28–29	26–37	25	-	21–25	-	20–33	Absent	~23

FIGURE 8. Potthastia longimanus (Kieffer, 1922) adult male (A–B), Lake Huron, Bush Bay, MI, USA. A. hypopygium, arrows indicate phallapodeme. B. hypopygium, aedeagal lobe (AL), superior volsella (SVo), median volsella (MVo), and inferior volsella (IVo), white arrows indicate the boundary of aedeagal lobe and superior volsella. Scale bars are 50 µm. FIGURE 9. Bayesian Interface (BI) and Neighbor-Joining (NJ) trees of Sympotthastia Pagast, 1947 species , and one outgroup Lappodiamesa omelkoi Makarchenko et Makarchenko, 2013 inferred from the COI nucleotide sequence data (658 bp). Numbers on branches represent the posterior probabilities for BI and bootstrap value for NJ, respectively (values <0.95/95 omitted). The gray bars on the right are clusters estimated using three molecular species delimitation methods: ABGD, ASAP, and K2P. In constructing Table 6 , we consulted the relevant literature that described the Sympotthastia species , and in doing so, we have also made corrections to the previous measurement records. Makarchenko (1985) provided an AR value of 3.48–3.69 for Sympotthastia takatensis (Tokunaga, 1936) , under Sympotthastia khorensis . Liu et al. (2016) indicated an AR value of 3.19–3.48 for this species. However, in their diagnosis and the key to adults of Sympotthastia they indicated the AR value of 3.19–4.26. Further, Makarchenko et al. (2022a) indicated an AR value of 2.76–2.81 for S. takatensis from the Republic of Khakassia , Russia . Dr. Wenbin Liu has kindly confirmed that their AR value of 3.19–4.26 is extracted from all S. takatensis reviewed and measured from Russia , Japan , and the Korean Peninsula. Dr. E. Makarchenko has also confirmed their value of 2.76–2.81 for S. takatensis from the Republic of Khakassia. Therefore, it seems this species has a large AR variation across its geographical range, so we considered an AR value of 2.8–4.3 for S. takatensis . This value falls within the AR range value for Sympotthastia huldeni Tuiskunen. Given the similarities in the shapes and sizes of anal point and AL + SVo of S. takatensis and S. huldeni along with their similar AR value, the only characteristic that separates them is the surface of S. takatensis median volsella bearing spinules and its apical edge being smooth whereas S. huldeni has a median volsella with bare surface and serrated apical edge. Dr. Liu has further provided us with a correct VR value of 0.83–0.87, for S. wuyiensis , which should replace the value of 3.29–3.44 previously provided by Liu et al. (2016) . Regarding the absence of values for some characteristics of Sympotthastia macrocera Serra-Tosio, 1973 in Table 6 , Serra-Tosio (1969) indicated that besides the ratio of palp length to head width, antennal ratio, and more temporal setae, S. macrocera is virtually the same as Sympotthastia spinifera Serra-Tosio, 1969 . At the time, he described the species as a variation of S. spinifera ; however, later in his 1973 work, Serra-Tosio erected the species based on discovering several other adults and immatures and the consistency of the distinctive characteristics.However, he did not provide any additional description or measurements. The total length and wing length measurements of this species in Table 6 are based on Doughman (1985) . There seems to be an inconsistency regarding the morphology terminologies used for the hypopygium of Sympotthastia . Liu et al. (2016) have indicated that what is referred to as superior volsella is, in fact, the aedeagal lobe. This terminology usage follows that of Hansen & Cook (1979) and Oliver (1989) , which refer to the structure as the aedeagal lobe or median aedeagal lobe. Hansen & Cook (1979) used the terminology proposed by Saether (1971) while discussing the difficulty of finding the correct homology of the external genitalia in insects and their correspondence to Chironomidae (see also Montagna et al. 2016b ). According to Saether (1971) , the aedeagal lobe is a membranous lobe bearing two dorsal sclerotized members that articulate anteriorly with the sternapodeme and converge posteriorly to nearly touch each other. Saether (1980) further described this lobe as an intromittent organ resulting from the combination of gonapophysis IX and surrounding phalli, which he suggested only truly exists in Buchonomyiinae . Saether’s (1980) work solidified the terminology of volsellas, especially in relation to the phallapodeme and aedeagal lobe. While defining these structures, their association with the aedeagal lobe can be extracted, with inferior volsella as the lateral lobe of the aedeagus, median volsella as the dorsal lobe of the aedeagus, and superior volsella as aedeagal blade. Further, Saether’s (1980) illustrations provided for species of Tanypodinae and Diamesinae male genitalia support this association where at least the superior and median volsellas are located in the aeadeagal region of the hypopygium attached to phallapodeme (i.e., surrounding phalli). Therefore, the definition of superior volsella proposed by Doughman (1985) and consequently used by Makarchenko (1994) is not necessarily incorrect. Doughman (1985) defined the superior volsella as an anteromedial portion of the phallapodeme that is apodemal and heavily sclerotized and the median volsella as a secondary, microsetigerous lobe of the phallapodeme. He defined the hirsute lobe, which is poorly developed, as inferior volsella ( Doughman 1985 ). From these, it is clear that what workers refer to as aedeagal lobe or superior volsella interchangeably (E. Makarchenko and B. Rossaro, pers. com. August 12, 2024 ) both have merits because the origin of these structures is the same. We can further find support in the description of the genitalia of the related families of Diptera , particularly Simuliidae . In the genitalia of adult male Simuliidae , the ventral plate of the aedeagus (superior volsella) is a large, heavily sclerotized platelike structure situated between and dorsal to gonopods with anterolateral corners being arm-like and median sclerite (median volsella) is usually moderately long, slender, flattened, and lying immediately dorsal to a ventral plate of the aedeagus ( Peterson 1981 ). What we observed in our specimens was that the posterior portion of what is called superior volsella by some workers and the aedeagal lobe by others appears much darker and indiscernibly separated from the much lighter anterior portion ( Figs. 5E & 6E ). This suggests that the posterior half should be regarded as the aedeagal lobe and the anterior half as the superior volsella so that the two are merged into one sclerite. By examining these sclerites in the closely related genus Potthastia , we can observe a clear separation of the aedeagal lobe from the superior volsella yet observe at the same time that they form the same plane ( Fig. 8 ). However, since the boundary of where the aedeagal lobe terminates and superior volsella begins may not be clearly observable in all Sympotthastia species , we suggest that the two be used in combination so that when referring to this sclerite, we use aedeagal lobe + superior volsella or AL + SVo.