Chilelimnophila Alexander, 1968

Krzemiński, Wiesław, Kania-Kłosok, Iwona, Arillo, Antonio, Kopeć, Katarzyna, Santos, Daubian & Soszyńska, Agnieszka, 2024, The evolutionary history and biogeographical distribution of the Mesozoic relic genus Chilelimnophila (Diptera, Limoniidae), Zoological Journal of the Linnean Society 202 (1) : -

Genus: Chilelimnophila Alexander, 1968

Type species: Limnophila lyra Alexander, 1952 (original designation)

Emended diagnosis: The genus is defined by the unique set of features that are not present together in other Limoniidae genera such as: antenna 11-segmented or 16-segmented, absence of crossvein r-r (R 2), presence of four medial veins that extend to the wing’s edge, and a distinct fusion of the first three flagellomeres in a male. The antennae of Chilelimnophila are distinguishable by the enlarged pedicel and a constriction in the first flagellomere.

Remarks: Since the genus was previously monotypic, we expanded its diagnosis to incorporate new features observed in fossil species, in comparison to the diagnosis provided by Ribeiro (2007).

Subgenus: Chilelimnophila (Ribeironia) Krzemiński, Kania-Kłosok et Arillo , subgen. nov.

Type species: Chilelimnophila (Ribeironia) amorimi Krzemiński, Kania-Kłosok et Arillo, sp. nov. ( Fig. 2).

Zoobank registration: urn:lsid:zoobank.org:act : FB3C6206- 010A-4D7D-8134-D78538144426

Diagnosis: Antenna 11-segmented; pedicel is small, barrelshaped, flagellomeres are short, oval and only slightly elongated, with short bristles no longer than the length of the segments on which they are located.

Etymology: The subgeneric name is dedicated to eminent entomologist Professor Guilherme Cunha Ribeiro from University de São Paulo, specialist on extant and extinct insects.

Chilelimnophila (Ribeironia) amorimi Krzemiński, Kania-Kłosok et Arillo, sp. nov.

( Fig. 2A–F)

Zoobank registration: urn:lsid:zoobank.org:act:7C7A1D3B-FE45-4FF5-BC5E-ADF0518F3A05

Diagnosis: Vein R 1 elongate, tip of R 1 positioned far beyond fork of R 2 + 3+4; sc-r situated before tip of Sc, approximately five times the distance from the tip of Sc; outer gonostylus narrow, elongate, strongly sclerotized, pointed, slightly curved to the internal part of hypopygium; inner gonostylus longer than half the length of the outer one.

Material examined: Holotype No. MCNA 9845 (male) housed in in the Museo de Ciencias Naturales de Álava , ( Vitoria-Gasteiz , Álava, Spain). Specimen well preserved, with partially destroyed legs and left antenna.

Horizon and locality: Lower Cretaceous (Upper Albian); Peñacerrada I amber site (Peñacerrada I, Moraza), eastern area of the Basque-Cantabrian Basin, Burgos, northern Spain.

Description: Body ( Fig. 2A, B) 1.80 mm long, dark brown, wings without colour pattern, pterostigma absent.

Head: Antenna ( Fig. 2C) 0.50 mm long, longer than head, longer than palpus; scape cylindrical, very elongate, approximately 4× as long as wide, pedicel ovoid, constricted at base, scape approximately twice as long as pedicel; first flagellomere ovoid, constricted at base; first flagellomere ovoid; flagellomeres 2‒11 elongate, approximately twice as long as wide, becoming more slender to the apex of antenna, first flagellomere only slightly longer and wider than second one, first flagellomere partially fused with the second one, last flagellomere equal in length to penultimate one; last flagellomere with two, rather short setae at apex, each other flagellomeres with four not very elongate setae, shorter than segments bearing them; maxillary palpus foursegmented, rather short, 0.3× the length of antenna, 0.20 mm long, palpomeres more or less cylindrical, not very elongate, longer than wide, rather narrow, third palpomere longer than second and fourth, slightly widened in midlength, second and fourth palpomere 2× as long as wide, third palpomere approximately 2.5× as long as wide; all palpomeres ( Fig. 2D) with few not very elongate setae, shorter than segments bearing them.

Thorax: Almost as long as high; wing 2.20 mm long, 0.80 mm wide ( Fig. 2F); h vein situated between the base of wing and fork of Mb and Cu; Sc ending on C before the level of bifurcation of Rs and tip of A 1; sc-r situated at approximately six of its length from the tip of Sc; Rs slightly curved, originating well proximally to the level of the tip of A 2, only slightly longer than R 2 + 3+4, R 1 reaches 0.3 the level of petiole, fork of R 2 + 3+4 situated just beyond the level of d-cell, R 3 approximately 0.6× the length of R 4; R 4 longer than R 2 + 3+4, longer than Rs; section of R 5 between its origin and point of contact with crossvein r-m straight; vein M 3 almost straight, approximately 1.5× the length of d-cell; d-cell 2× as long as wide; crossvein m-cu shorter than d-cell; A 1 almost straight, tip of A 1 situated beyond fork of Mb; A 2 slightly curved; tip of A 2 just before the level of m-cu; tip of A 2 situated just before the level of fork of Mb, beyond the level of fork of Rb; haltere: stem narrow, elongate, slightly longer than knob; tibial spurs rather thick, not very elongate, shorter than setae which occur on tibia.

Abdomen: Male terminalia ( Fig. 2E)—hypopygium 0.20 mm long, elongate and narrow, gonocoxite approximately 3× as long as wide, tip of gonocoxite and outer gonostylus strongly sclerotized, almost black, outer gonostylus narrow, pointed, slightly curved to internal part of hypopygium, outer gonostylus rather slender but lobe-shaped, very pale, rounded at apex.

Etymology: The specific name is dedicated to eminent entomologist and paleontologist, Professor Dalton de Souza Amorim from the Departamento de Biologia (FFCLRP), Universidade de Sao Paulo, Brazil.

Remarks: The antenna of C. (R.) amorimi subgen. et sp. nov. is reduced and 11-segmented, while other species of Chilelimnophila are characterized by 16-segmented antenna.

Subgenus: Chilelimnophila (Chilelimnophila) Alexander, 1968 .

Type species: Limnophila lyra Alexander, 1952 (original designation).

Emended diagnosis: Antenna 16-segmented; pedicel very large, oval; first three flagellomeres short, broad, while other flagellomeres narrow, very elongated, with long bristles some of which are much longer than the flagellomere on which they are located.

Remarks: Diagnosis emended in comparison to the diagnosis provided by Ribeiro (2007).

Chilelimnophila (Chilelimnophila) wangi Krzemiński, Kania-Kłosok et Arillo, sp. nov.

( Fig. 3A–E)

Zoobank registration: urn:lsid:zoobank.org:act : 665347C3- 157F-4E27-88DE-4B2BAB670023

Diagnosis: Vein R 1 elongate, tip of R 1 positioned far beyond fork of R 2 + 3+4; sc-r situated before tip of Sc, approximately nine times the distance from the tip of Sc; outer gonostylus narrow, elongate, slender, dark at the tip; inner gonostylus short and massive, reaching approximately half the length of gonocoxite, slightly curved to the internal part of hypopygium, sharpened at apex.

Material examined: Holotype No. MP/4449 (male), coll. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland ( ISEA PAS). Specimen well preserved, with partially destroyed legs and wing.

Horizon and locality: Lowermost Cenomanian , Hukawng Valley , northern Myanmar. The mining is done at a hill named Noije Bum, near Tanai Village (26 ° 21′ 33.41′′ N, 96 ° 43′ 11.88′′ E) .

Description: Body ( Fig. 3D) 2.69 mm long, brown, wings without colour pattern, pterostigma absent.

Head: Antenna ( Fig. 3A) 0.85 mm long, longer than head, longer than palpus; scape cylindrical, rather short, only slightly longer than pedicel; approximately 2× as long as wide, pedicel ovoid, constricted at base, scape approximately 1.5× as long as pedicel; first flagellomere fused with second and third; flagellomeres 5‒14 elongate, tiny, approximately 4–5× as long as wide, become more slender to the apex of antenna, first flagellomere longer and wider than second one, second and third flagellomeres very short, fused with the first one; last flagellomere shorter than penultimate one; each flagellomeres with three to five elongate setae, longer than segments bearing them; maxillary palpus foursegmented ( Fig. 3B), short, 0.2× the length of antenna, 0.21 mm long, palpomeres rather ovoid, short, only slightly longer than wide, third palpomere comparable length to the second, all palpomeres slightly widened in midlength; all palpomeres with few setae, shorter than segments bearing them.

Thorax: Almost as long as high ( Fig. 3D); wing 2.94 mm long, 0.84mm wide ( Fig. 3E); h vein situated between the base of wing and fork of Mb and Cu; Sc ending on C at the level of bifurcation of Rs and tip of A 1; sc-r situated at approximately eight of its length from the tip of Sc; Rs slightly curved, originating well proximally to the level of the tip of A 2, only slightly longer than R 2 + 3+4, R 1 reaches almost half the length of petiole; fork of R 2 + 3+4 situated just beyond the level of d-cell, R 3 approximately 0.6× the length of R 4; R 4 longer than R 2 + 3+4, longer than Rs; section of R 5 curved between its origin and point of contact with crossvein r-m; vein M 3 almost straight, approximately 1.5× the length of d-cell; d-cell 3× as long as wide; crossvein m-cu shorter than d-cell; A 1 slightly curved at the tip, tip of A 1 situated beyond fork of Mb; A 2 slightly curved; tip of A 2 just before the level of m-cu, just before the level of fork of Mb, beyond the level of fork of Rb; haltere: stem narrow, elongate, slightly longer than knob; tibial spurs rather thick, not very elongate, shorter than setae which occur on tibia.

Abdomen: Male terminalia ( Fig. 3E)—hypopygium 0.55 mm long, elongate and narrow, gonocoxite approximately 4× as long as wide, tip of gonocoxite and outer gonostylus slightly sclerotized, brown, outer gonostylus narrow, pointed, slightly curved to internal part of hypopygium, inner gonostylus rather slender but lobe-shaped, very pale, rounded at apex.

Etymology: The specific name is dedicated to eminent palaeontologist Professor Bo Wang from the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China.

Chilelimnophila (Chilelimnophila) parva Krzemiński, Kania-Kłosok et Arillo, sp. nov.

( Fig. 4A–E)

Zoobank registration: urn:lsid:zoobank.org:act:8B72FF8D-EA83-4A2F-9085-7F5892A658BD

Diagnosis: Tip of vein R 1 approximately at fork of R 2 + 3+4; sc-r situated before tip of Sc, approximately two times the distance from the tip of Sc; gonocoxite very elongate, approximately 4× as long as wide, widened in 0.3× of its length and folded to the internal part of hypopygium; outer gonostylus narrow, slightly sclerotized, elongate, longer than half the length of gonocoxite, slightly curved to the internal part of hypopygium; inner gonostylus short and massive, round at apex, shorter than half the length of outer gonostylus, slightly curved to the internal part of hypopygium.

Material examined: Holotype No. MP/4509 (male), coll. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland ( ISEA PAS). Specimen well preserved.

Horizon and locality: Lowermost Cenomanian , Hukawng Valley , northern Myanmar. The mining is done at a hill named Noije Bum, near Tanai Village (26° 21′ 33.41′′ N, 96° 43′ 11.88′′ E) .

Description: Body ( Fig. 4B) 2.56 mm long, brown, wings without colour pattern, pterostigma absent.

Head: Antenna ( Fig. 4A) 1.03 mm long, longer than head, longer than palpus; scape cylindrical, slightly elongate, comparative length to pedicel; approximately 2× as long as wide, pedicel ovoid, constricted at base, first flagellomere fused with second and third; flagellomeres 4–16 elongate, tiny, approximately 2–3× as long as wide, become more slender to the apex of antenna, first flagellomere as long as wide, second and third flagellomeres rather short, fused with the first one; last flagellomere as long as penultimate one; each flagellomeres with two elongate setae, longer than segments bearing them; maxillary palpus foursegmented, short, 0.2× the length of antenna, 0.24 mm long, palpomeres rather ovoid, short, only slightly longer than wide, third palpomere comparable length to the second, all palpomeres slightly widened in the mid length; all palpomeres with few not very elongate setae, as long or shorter than segments bearing them.

Thorax ( Fig. 4B): Almost as long as high; wing 3.40 mm long, 1.13 mm wide ( Fig. 4B, E); h vein situated between the base of wing and fork of Mb and Cu; Sc ending on C before the level of bifurcation of Rs and tip of A 1; sc-r situated at approximately two of its length from the tip of Sc; Rs slightly curved, originating well proximally to the level of the tip of A 2, only slightly longer than R 2 + 3+4, R 1 reach 0.25× the length of petiole; fork of R 2 + 3+4 situated just beyond the level of d-cell, R 3 approximately 0.6× the length of R 4; R 4 longer than R 2 + 3+4, longer than Rs; section of R 5 between its origin and point of contact with crossvein r-m almost straight; vein M 3 almost straight, approximately 2× the length of d-cell; d-cell 2× as long as wide; crossvein m-cu shorter than d-cell; A 1 almost straight, tip of A 1 situated beyond fork of Mb; A 2 slightly curved; tip of A 2 just before the level of m-cu, just before the level of fork of Mb, beyond the level of fork of Rb; haltere: stem narrow, elongate, slightly longer than knob.

Abdomen: Male terminalia ( Fig. 4D)—hypopygium 0.53 mm long, elongate and narrow, gonocoxite approximately 4× as long as wide, tip of gonocoxite and outer gonostylus slightly sclerotized, brown, outer gonostylus narrow, pointed, slightly curved to internal part of hypopygium, inner gonostylus rather slender but lobe-shaped.

Etymology: The specific name is derived from ‘ parva ’ (Latin)—small.

Comparison: The main differences between Recent and Cretaceous species of Chilelimnophila are the morphology of the hypopygium ( Figs 2–4) and wing venation ( Fig. 5). In these newly described Cretaceous species, the gonocoxite is narrow and elongate, in C. (R.) amorimi subgen. et sp. nov. and C. (C.) wangi sp. nov. the coxite is approximately 3× as long as wide, in C. (C.) parva sp. nov. it is approximately 4× as long as wide, while in C. (C.) lyra the gonocoxite is massive and wide, but relatively shorter, at most 2× as long as wide. In C. (R.) amorimi subgen. et sp. nov. the tip of the gonocoxite and outer gonostylus are strongly sclerotized, almost black, while in C. (C.) lyra they are only slightly sclerotized and pale. In C. (C.) parva sp. nov. the gonocoxite is folded inwards in 0.3× of its length, the outer gonostylus is longer than half the length of the gonocoxite, the inner gonostylus is rounded at its apex, and shorter than half the length of outer gonostylus. In comparison, in C. (C.) wangi sp. nov. the outer gonostylus is shorter than half the length of the gonocoxite, the inner gonostylus is sharpened at the apex, longer than half the length of the outer gonostylus. Moreover, in C. (C.) lyra vein Sc ends on C at the level of bifurcation of Rs and tip of A 1 while in C. (R.) amorimi subgen. et sp. nov. it ends before, and in C. (C.) wangi sp. nov it ends at the level of bifurcation of Rs and before the level of tip of A 1. In C. (R.) amorimi subgen. et sp. nov. and C. (C.) wangi sp. nov. the tip of A 1 is positioned even beyond half the level of the d-cell, while in C. (C.) lyra the tip ends before its level. Position of sc-r in these three species is different also, in Cretaceous species C. (R.) amorimi subgen. et sp. nov. and C. (C.) wangi sp. nov. crossvein sc-r is situated far before the tip of Sc and before half the length of Rs, while in C. (C.) lyra even if the position of sc-r is variable, this vein occurs more or less near the tip of Sc and the fork of Rs. Rs in C. (C.) lyra is elongate, while in Cretaceous species it is rather short. Crossvein m-cu in C. (C.) lyra is situated distinctly beyond half the length of the d-cell, while in newly described species m-cu occurs before half the length of the d-cell. In addition, M 1 and M 2 are quite short in C. (C.) lyra in relation to the length of its petiole, the petiole is approximately 3× longer than M 1 or M 2, in C. (R.) amorimi subgen. et sp. nov. Cell M 1 is longer than its petiole, while the petiole and M 2 are of comparable length. In C. (C.) lyra the basal section of R 5 between its origin and point of contact with r-m is curved, in C. (R.) amorimi subgen. et sp. nov. this is straight and perpendicular to fork of Rs, while in C. (C.) wangi sp. nov. the basal section of R 5 is slightly curved and shifted to the apex of wing, but not so far as in C. (C.) lyra .

Key to species of Chilelimnophila

1. Antenna 11-segmented; r-m inserted distal to the level of fork of vein Mb ......................................................................................... ....................................................................................................................... Chilelimnophila (Ribeironia) amorimi sp. nov. ( Fig. 2)

- Antenna 16-segmented; r-m inserted proximal to the level of fork of Mb......................................................................................(2)

2. Crossvein m-cu positioned beyond half the length of d-cell; M 1 short, shorter than half the length of petiole; gonocoxite at most 2× as long as wide...................................................................................... Chilelimnophila (Chilelimnophila) lyra ( Fig. 5M)

- Crossvein m-cu positioned before half the length of d-cell; M 1 longer than half the length of petiole; gonocoxite longer than twice its width .............................................................................................................................................................................................(3)

3. Gonocoxite elongate, approximately 3× as long as wide, almost straight; outer gonostylus shorter than half the length of gonocoxite, inner gonostylus sharpened at apex, longer than half the length of outer gonostylus ................................................. .................................................................................................................. Chilelimnophila (Chilelimnophila) wangi sp. nov. ( Fig. 3)

- Gonocoxite elongate, approximately 4× as long as wide, folded inwards in 0.3× of its length; outer of gonostylus longer than half the length of gonocoxite, inner gonostylus round at apex, shorter than half the length of outer gonostylus ........................ .................................................................................................................. Chilelimnophila (Chilelimnophila) parva sp. nov. ( Fig. 4)

Phylogenetic analysis

The characters we used are defined and listed below (the number of steps of each character for k = 3 is shown within square brackets). The data matrix is presented in Table 1. The parsimony analysis with implied weights resulted in a single most-parsimonious tree shown in Figure 6 which retains the same topology of the EW. The most-parsimonious trees have 39 steps, consistency index (CI) = 0.84 and retention index (RI) = 0.91. The synapomorphies of each clade are shown in Figure 6.

0: Length of rostrum: 0—shorter than the remainder of the head; 1—as long as the remainder of the head; 2—at least two times longer than the remainder of the head. [2]

1: Width of pedicel: 0—similar in width than the first flagellomer; 1—at least two times wider than the first flagellomere. [1]

2 Shape of the flagellomeres: 0—cylindrical, at least three times longer than wide; 1—ovoid or subquadrate. [1]

3 Aspect of flagellomeres: 0—with the distal portion lighter and with some desclerotization; 1—uniform. [1]

4 Base of the first flagellomere: 0—wide; 1—with a constriction. [1]

5 Fusion of the firsts flagellomeres (in males): 0— flagellomeres completely separated; 1—firsts flagellomeres partially or totally fused. [1]

6 Stage of fusion of flagellemores:0—only a partially fusion of the flagellomeres 1 and 2; 1—fusion of flagellomeres 1-3; 2—fusion of flagellomeres 1-4. [2]

7 Proportions of the second flagellomere: 0—longer than wide; 1—as long as wide, subquadrate in shape. [1]

8 Proportions of the third flagellomere: 0—longer than wide; 1—as long as wide, subquadrate in shape. [1]

9 Length of coxa 2: 0—shorter than katepisternum; 1— similar or longer than katepisternum. [1]

10 Length of sc-r: 0—similar in length than the distal section of Sc; 1—at least twice the length of the distal section of Sc; 2—short, not surpassing the third of the length of the distal section of Sc. [2]

11 Vein R

2

: 0—present; 1—absent. [1]

12 Level of insertion of R 2: 0—in the middle of discal cell; 1—more distal than discal cell. [2]

13 Directions of the veins R 3 and R 4: 0—divergent; 1—almost straight and running parallel. [2]

14 Level of insertion of r-m: 0—at same level of Mb fork; 1—more proximal than Mb fork. [3]

15 Vein petiole: 0—forked on M 1 and M 2; 1—unbranched. [1]

16 Length of petiole: 0—tiny, many times shorter than M 1; 1—shorter or similar in length to M 1; 2—longer than M 1. [2]

17 Length of m-m: 0—half or shorter the length of the basal section of M 3 (from the fork of M 3 + 4 until the contact with m-m); 1—as long as the basal section of M 3. [1]

18 Shape of the basal section of M 3: 0—straight; 1— curved. [1]

0123456789012345678901234567 Paralimnophila skusei 000000-000000000100000000000 Ctenolimnophila 001100-000001011-01100011010 (Campbellomyia) paulistae

Ctenolimnophila 001100-00?000001-01100011010 (Ctenolimnophila) bivena

Edwardsomyia chiloensis 200000-001001100000100010001 Mesolimnophila lutea 101100-000001100100100011110 Mesolimnophila hirsutipes 101100-00?001100100100011110 Tonnoirella gemella 200000-001101100000100010001 Tonnoirella marmoripennis 200000-001101100000100010001 Tinemyia margaritifera 200000-000001110100100110001 Zelandomyia armigera 001100-00001-000210101011000 Zelandomyia pallidula 001100-00001-000210101011000 Chilelimnophila (C.) lyra 011111211021-010200100011000 Chilelimnophila (R.) 011111010?21-000200100111000 amorimi Ǫ. nov.

Chilelimnophila (C.) 011111111021-010200110111200 wangi sp. nov.

Chilelimnophila (C.) 011111111021-010200110111200 parva sp. nov.

19 Level of insertion of m-cu: 0—at the fork of Mb; 1—at the discal cell (vein M

3 + 4

). [1]

20 Gonocoxite; 0—stout, no more than three times longer than wide; 1—slender, at least 4 times longer than wide. [1]

21 Gonocoxite apodema; 0—reduced or absent; 1— strongly chitinized. [1]

22 Position of gonostylus: 0—terminal; 1—subterminal. [3]

23 Outer branch of gonostylus: 0—bifid; 1—unbranched. [1]

24 Length of the outer branch of gonostylus: 0—shorter than the inner branch; 1—similar or longer than the inner branch. [1]

25 Curvature of the outer branch of gonostylus: 0—only with the distal part curved; 1—straight; 2—at least the last third of the branch curved. [2]

26 Dorsal margin of the outer branch of gonostylus: 0— smooth; 1—serrated. [1]

27 Long setae in the outer branch of gonostylus: 0—absent; 1—present. [1]