Pilielmis nheengatu, Pinedo-Garcia & Polizei & Hamada & Nascimento, 2024

Pinedo-Garcia, Raul Bismarck, Polizei, Thiago Tadeu Silva, Hamada, Neusa & Nascimento, Jeane Marcelle Cavalcante Do, 2024, Systematics of Pilielmis Hinton, 1971 (Coleoptera: Dryopoidea: Elmidae), Zootaxa 5421 (1), pp. 1-68 : 54-65

Pilielmis nheengatu sp. nov.

( Figs 37–40 )

Diagnosis. Body subparallel; general coloration black; antennae bicolor, antennomeres I, II yellow, antennomeres III–XI dark brown; pronotum with a brown mark on anterior 1/3; elytra with yellowish-brown marks (a pair of subrectangular marks on humeral region; two pairs of elongated, sublateral marks approximately on apical 3/5 and 4/5, and a mark on the apex along the elytral suture). Male with phallobase 1.5x longer than wide; parameres reaching approximately half the length of the penis; penis approximately 3x longer than wide, with a constriction on basal 1/3; basolateral process 2x longer than wide.

Description. Male. Length: mean = 1.5 mm; width: mean = 0.65 mm (n = 2); Body subparallel ( Fig. 37A ); general coloration black, legs reddish-brown, pronotum and elytra with yellowish-brown marks.

Head. Antennae bicolor, antennomeres I–II yellow, antennomeres III–XI brown ( Fig. 37D ). Labrum ( Fig. 38A ) subrectangular, 1.75x longer than wide, with lateral margins rounded; anterolateral margins with long setae; dorsal surface with short, sparse setae. Mandibles ( Fig. 38C, D ): lateral lobe rounded; molar area with a groove at base; posterolateral angle slightly rounded; condyle semicircular, wider than long. Maxilla ( Fig. 38B ): cardo subtriangular; stipe subtriangular; maxillary palpomere IV elongated, as long as the two previous one combined, with apex rounded; galea with apical section subtriangular, 2x longer than wide, lateral margin and apex rounded, with long, thin setae; lacinia subrectangular, 2x longer than wide, with long setae on inner lateral margin. Labium ( Fig. 38E ): postmentum subtriangular, wider than long; prementum subrectangular, base and apex convex, 5x wider than long; ligula subrectangular, 2x wider than long, anterolateral regions laterally projected, with short spines; gula subrectangular, 4x wider than long, lateral margins straight, anterolateral angles sharp; labial palpomere III with rounded apex.

Thorax. Pronotum ( Fig. 37A ) slightly wider (0.45 mm) than long (0.41 mm); with short, golden setae; lateral margins sinuous, with crenulations; anterolateral angles acute; posterolateral angles narrow; anterior region narrower than posterior region; with a brown mark on the anterior 1/3; with a pair of straight, strongly raised sublateral carinae; longitudinal impression on the disc deep; with a carina on the anterior half. Elytra ( Fig. 37A ) subparallel, 1.5x longer than wide (length = 0.98 mm; maximum width = 0.65 mm); apex rounded; with yellowish-brown marks (a pair of subrectangular marks on humeral region; two pairs of elongated, sublateral marks approximately on apical 3/5 and 4/5, and a mark at the apex along the elytral suture). Scutellum subtriangular. Metathoracic wings present ( Fig. 41E ). Prosternum ( Fig. 37B ) 0.4x longer than wide, anterior margin concave, anterolateral angles acute. Prosternal process ( Fig. 37B ) approximately as long (0.08 mm) as wide (0.11 mm). Mesoventrite ( Fig. 37B ) wider (0.19 mm) than long (0.06 mm); posterior margin concave. Metaventrite ( Fig. 37B ): lateral margins covered by tomentum; discrimen conspicuous on basal 3/4; disc flat; surface with few, short setae. Hind legs ( Fig. 37B, C ). Femora 3.7x longer than wide. Tibiae 6.2x longer than wide. Apical tarsomere longer than the basal ones combined. Claws large and slender, with a tooth near base.

Abdomen. Ventrites I–IV without tomentum on disc; intercoxal process of ventrite I depressed; ventrite V with golden setae ( Fig. 37B ). Tergite VIII ( Fig. 39C ) almost as long as wide, posterior margin rounded; surface covered by spines and microspines. Sternite VIII ( Fig. 39D ): 3.5x wider than long; surface with setae near apex; anterior median projection 2x longer than the sternite body length. Segment IX ( Fig. 39B ) subtriangular, with rounded apex; anterior margin with sensory pores. Genitalia ( Fig. 39A ). Phallobase 1.5x longer than wide, approximately the same length as the penis. Parameres reaching approximately half the length of the penis, base wider than the apex; apex rounded. Penis approximately 3x longer than wide, with a constriction on basal 1/3; apical region rounded; lateral margin sinuous; surface with sensory micropores throughout the area; basolateral process 2x longer than wide.

Female. Externally similar to male. Length = 1.64 mm; width = 0.67 mm (n = 1). Tergite VIII ( Fig. 40B ) almost as long as wide, posterior margin slightly acuminated, with long setae; surface with microspines and short setae. Sternite VIII ( Fig. 40C ) 2x wider than long; anterior median projection 2x longer than the sternite body length; posterior margin with long setae, surface with short setae and microspines. Ovipositor ( Fig. 40A ): valvifer posteriorly dilated; 2x longer than the coxites. Apical coxite 2.4x longer than wide, 1.5x longer than the basal one. Stylus slightly curved, measuring 1/4 the length of the apical coxite; apex with three short setae.

Etymology. The specific epithet nheengatu is in honor of the Nheengatu language that is the most spoken by the indigenous peoples that inhabitants the municipality of São Gabriel da Cachoeria (type locality of this species). This name is used in apposition.

Comparative notes. Pilielmis nheengatu sp. nov. can be distinguished from any other species of Pilielmis based on the unique pattern of coloration, mainly by the blackish general coloration. In addition, the shape of the penis of this species is unique, with a constriction on its basal 1/3. Among the species currently known for Pilielmis , the only one with a blackish general coloration is P. shepardi ; however, the shape of the body and the elytral marks can easily differentiate between these species. The new species has a subparallel body, and the elytra has elongated marks on the apical 3/5 and 4/5. On the other hand, P. shepardi has a suboval body and the elytra has only one pair of suboval marks on the central region.

Distribution. Brazil (Amazonas) ( Fig. 1 ).

Type material. Holotype. Male. Brazil, Amazonas , São Gabriel da Cachoeira , Pari-Cachoeira, Samaúma stream, 17-20.iii.2016, 00°15′34.7′′ S, 69°47′15.3′′ W, Hamada & Nascimento leg., ( INPA —body in alcohol; mouthparts and genitalia on slide) . Paratypes: same data as holotype (1♂ and 1♀ in INPA; 1♂ in MZUSP) ( INPA —body in alcohol; mouthparts, wing, and genitalia on slide) .

Identification key for adults of Pilielmis species

1. Body suboval ( Figs 3A , 27A ); metathoracic wings absent..................................................... 2

1’. Body subparallel ( Figs 7A , 19A , 23A ); metathoracic wings present............................................. 3

2. General coloration dark brown ( Fig. 3A ); metaventrite disc with a pair of well-developed sublateral carinae ( Fig. 3B ); ventrite V and sternite VIII not fused; males with parameres reaching half the length of the penis, with a long seta at the apex; penis with rounded apex ( Fig. 5A )..................................................................... P. murcia

2’. General coloration black ( Fig. 27A ); metaventrite disc without carinae ( Fig. 27B ); ventrite V and sternite VIII fused; males with parameres reaching 3/5 the length of the penis, without setae at the apex; penis with subtriangular apex ( Fig. 29A )................................................................................................... P. shepardi

3. Antennae with uniform coloration ( Figs 3A , 19A , 23A , 27A ).................................................. 4

3’. Antennae bicolor ( Figs 7A , 11A , 15A , 31A )............................................................... 7

4. Elytra with apex rounded, without projections ( Figs 19A , 23A , 37A )............................................ 5

4’. Elytra with apex with a pair of short projections ( Fig. 34A )..................................... P. muricula sp. nov.

5. General coloration yellowish-brown or brown; humeral region without marks ( Figs 19A , 23A )....................... 6

5’. General coloration blackish; humeral region with yellowish marks ( Fig. 37A ).................... P. nheengatu sp. nov.

6. Prosternal process longer than wide ( Fig. 19B, C ); femora with uniform coloration ( Fig. 19C ); male with phallobase 2.3x longer than wide, penis 3x longer than wide, basolateral process 3.5x longer than wide ( Fig. 21A )..................... P. clita

6’. Prosternal process as long as wide ( Fig. 23B ); femora with apical region darker ( Fig. 21B, C ); male with phallobase 1.8x longer than wide, penis 4x longer than wide, basolateral process 6x longer than wide ( Fig. 25A )..................... P. abdera

7. Posterior region of pronotum with a pair of black marks ( Figs 9A , 31A )......................................... 8

7’. Posterior region of pronotum without a pair of black marks ( Figs 11A , 15A )...................................... 9

8. Elytra yellowish-brown, with a pair of black marks ( Fig. 7A ); metaventrite with a pair of short carinae at the base of the discrimen; male genitalia with parameres reaching 3/5 the length of the penis; lateral margin of paramere strongly crenulated; penis 2.5x longer than wide, lateral margins with a strong constriction ( Fig. 9A ).............................. P. sara

8’. Elytra black, with yellow marks ( Fig. 31A ); metaventrite without a pair of short carinae at the base of the discrimen; male genitalia with parameres reaching 3/4 the length of the penis; lateral margin of paramere smooth; penis 4x longer than wide, lateral margins without strong constriction ( Fig. 33A ).......................................... P. baniwa sp. nov.

9. Metatrochanter with a gibbosity on the apical region ( Fig. 11B ); females with a tuft of long, thin setae at apex of the metatibia; males with an excavation on the metatibia; phallobase 1.5x longer than wide, lateral margins with a constriction on medial region; penis 2.7x longer than wide; basolateral process 3x longer than wide ( Fig. 13A )........................ P. halia

9’. Metatrochanter without gibbosity on the apical region ( Fig. 15B ); females without a tuft of long, thin setae at apex of the metatibia; males without excavation on the metatibia; phallobase 2.3x longer than wide, lateral margins straight, without constriction; penis 4.4x longer than wide; basolateral process 7.2x longer than wide ( Fig. 17A )................ P. apama

Phylogeny

The phylogenetic study resulting in a matrix (Appendix I) with 42 characters listed below. For each character its length (L), consistency index (CI) and retention index (RI) is presented.

Character list

1. Body, length versus width, ratio (L = 1; CI = 100; RI = 100):

(0) up to 2x longer than wide (suboval)

(1)> 2x longer than wide (subparallel)

2. Antennae, coloration pattern (L = 2; CI = 50; RI = 85):

(0) unicolor

(1) bicolor

3. Labrum, shape (L = 1; CI = 100; RI = 100):

(0) suboval

(1) subrectangular

4. Maxilla, apex of galea, shape (L = 5; CI = 20; RI = 20):

(0) rounded

(1) truncated

5. Maxilla, apex of palpomere IV, shape (L = 3; CI = 33; RI = 50):

(0) rounded

(1) truncated

6. Labium, apex of palpomere III, shape (L = 1; CI = 100; RI = 100):

(0) rounded

(1) truncated

7. Pronotum, longitudinal impression on disc, development (L = 5; CI = 20; RI = 33): (0) deep

(1) shallow

8. Pronotum, sublateral carinae, shape (L = 5; CI = 40; RI = 40):

(0) straight

(1) sinuous

(2) convex

9. Pronotum, sublateral carinae, development (L = 2; CI = 50; RI = 66):

(0) complete

(1) incomplete

10. Pronotum, sublateral carinae, elevation, development (L = 2; CI = 50; RI = 75): (0) strongly elevated

(1) slightly elevated

11. Pronotum, posterior half, carinae (L = 4; CI = 25; RI = 25):

(0) absent

(1) present

12. Pronotum, posterior region, pair of blackish marks (L = 2; CI = 50; RI = 0):

(0) absent

(1) present

13. Pronotum, anterolateral angles, shape (L = 3; CI = 33; RI = 66):

(0) acute

(1) rounded

14. Pronotum, lateral margin, shape (L = 3; CI = 33; RI = 0):

(0) convex

(1) sinuous

15. Pronotum, lateral margin, crenulation (L = 1; CI = 100; RI = 100):

(0) present

(1) absent

16. Elytra, basal carina on interval IV (L = 1; CI = 100; RI = 100):

(0) absent

(1) present

17. Elytra, sublateral carina on interval VIII (L = 2; CI = 50; RI = 75):

(0) present

(1) absent

18. Elytra, punctuation, development (L = 4; CI = 25; RI = 40):

(0) weak

(1) strong

19. Elytra, apex, shape (L = 2; CI = 100; RI = 100):

(0) rounded

(1) projected

(2) truncated

20. Elytra, scutellum, length versus width, ratio (L = 4; CI = 50; RI = 0):

(0) as wide as long

(1) longer than wide

(2) wider than long

21. Thorax, metathoracic wings (L = 2; CI = 50; RI = 50):

(0) absent

(1) present

22. Thorax, prosternum, length versus width, ratio (L = 4; CI = 25; RI = 50):

(0) ≥ 0.5x longer than wide

(1) <0.5x longer than wide

23. Thorax, prosternal process, posterior margin, shape (L = 2; CI = 100; Ri = 100): (0) rounded

(1) truncated

(2) bilobed

24. Thorax, metaventrite, carinae (L = 3; CI = 33; RI = 0):

(0) absent

(1) present

25. Thorax, metaventrite, oval depression (L = 1; CI = 100; RI = 100):

(0) absent

(1) present

26. Hind leg, femora, length versus width, ratio (L = 4; CI = 25; RI = 50):

(0) ≤ 3.9 longer than wide

(1) ≥ 4 longer than wide

27. Hind leg, tibiae, length versus width, ratio (L = 1; CI = 100; Ri = 100):

(0) ≤ 8x longer than wide

(1)> than 8x longer than wide

28. Hind leg, tibiae, row of spines (L = 1; CI = 100; RI = 100):

(0) absent

(1) present

29. Legs, claw, basal tooth (L = 2; CI = 50; RI = 66):

(0) absent

(1) present

30. Abdomen, ventrite I, anterior margin, ratio (L = 2; CI = 50; RI = 0):

(0) absent

(1) present

31. Abdomen, ventrite II, tomentum on disc (L = 4; CI = 25; RI = 25):

(0) absent (Fig. 74A)

(1) present (Fig. 74B)

32. Abdomen, ventrite IV, tomentum (L = 4; CI = 25; RI = 0):

(0) absent

(1) present

33. Abdomen ♂, tergite VIII, posterior margin, shape (L = 2; CI = 50; RI = 0) :

(0) rounded

(1) truncated

34. Genitalia ♂, penis length versus parameter length, ratio (L = 3; CI = 33; RI = 0):

(0) penis up to 2x longer than parameres

(1) penis longer than 2x the paramere length

35. Genitalia ♂, penis, apical region, shape (L = 4; CI = 25; RI = 50):

(0) subtriangular

(1) rounded

36. Genitalia ♂, penis, length versus width, ratio (L = 6; CI = 33; RI = 20):

(0) ≤ 2.9x longer than wide

(1) 3 to 3.9x longer than wide

(2) ≥ 4x longer than wide

37. Genitalia ♂, penis, lateral margins, shape (L = 1; CI = 100; RI = 100):

(0) straight

(1) sinuous

38. Genitalia ♂, phallobase, length versus width, ratio (L = 4; CI = 25; RI = 0):

(0) ≤ 1.9x longer than wide

(1) ≥ 2x longer than wide

39. Genitalia ♀, ovipositor, inner margin of apical coxite, row of setae (L = 1; CI = 100; RI = 100): (0) absent

(1) present

40. Genitalia ♀, ovipositor, apical coxite, external margin of distal region, projection (L = 0):

(0) absent

(1) present

41. Genitalia ♀, length of apical coxite versus length of basal coxite, ratio (L = 2; CI = 100; RI = 100): (0) up to 0.9x (apical coxite shorter than basal coxite)

(1) apical coxite 1 to 1.9x longer than basal coxite

(2) apical coxite ≥ 2x longer than basal coxite

42. Genitalia ♀, apical coxite, length versus width, ratio (L = 7; CI = 28; RI = 16):

(0) ≤ 1.9x longer than wide

(1) 2 to 2.9x longer than wide

(2) ≥ 3x longer than wide

The cladistic analysis under equal weights recovered the six most parsimonious trees with length (L) = 111, consistency index (CI) = 43 and retention index (RI) = 52. On the other hand, the analyses under implied weights resulted in a single tree, and the same topology was achieved in all K -values tested. The phylogenetic results presented below were based in the most parsimonious tree resulting from the analysis under K = 1 ( Fig. 42 ). The strict consensus of the six most parsimonious trees from the analysis under equal weights is shown in the Figure 43 .

Pilielmis was recovered as a monophyletic group (node “A”) supported by a synapomorphy: hind tibia ≤ 8x longer than wide (character 27 [state 0]). In addition, three non-unique synapomorphies support the monophyly of this genus: pronotum with incomplete sublateral carinae (9[1]), absence of sublateral carina on elytral interval VIII (17[0]), and tarsal claws with a basal tooth (29 [1]). Among the genera selected as outgroups in the analysis, Hexacylloepus appeared as sister genus of Pilielmis , and this relationship was supported by five synapomorphies, among them: the presence of a basal carina on elytral interval IV (16[1]) and females with the inner margin of the apical coxite with a row of setae (39[1]).

Within Pilielmis , some clades were recovered. In the first clade (node “B”), P. murcia appeared related to P. shepardi and these species share: body suboval (1[0]) and metathoracic wings absent (21[0]). The second recovered clade, represented by node “C”, is composed by the remaining species of Pilielmis . This clade is sustained by one non-unique synapomorphy: pronotum with anterolateral angles rounded (13[1]) (with reversion in P. clita (13[0])).

In the clade represented by the node “C”, two other clusters were recovered. The first (node “D”) is composed by P. nheengatu sp. nov. ( P. muricula sp. nov. ( P. clita + P. abdera )) and was supported by two non-unique synapomorphies: pronotum with sublateral carinae strongly elevated (10[0]) and with a carina on the basal half (11[1]) (with reversion in P. clita (11[0])). Pilielmis muricula sp. nov. appeared as a sister group of P. clita + P. Abdera (node “E”); this clade was supported by palpomere IV with apex truncated (5[1]). The clade composed by P. clita + P. abdera (node “F”) was supported by two synapomorphies and two non-unique synapomorphies: labial palpomere III with apex truncated (6[1]), metaventrite with oval depression (25[1]), apical palpomere of the galea with apex truncated (4[1]) and pronotum with a shallow longitudinal impression on disc (7[1]).

The node “G”, formed by P. baniwa sp. nov. ( P. apama ( P. sara + P. halia )), was supported by a non-unique synapomorphy: the presence of antennae bicolor (2[1]). Within this clade, P. apama appeared as a sister group of P. sara + P. halia (node “H”) and this relationship was supported by the prosternum <0.5x longer than wide (22[1]). Finally, P. sara was recovered as a sister species of P. halia supported by one synapomorphy (labrum suboval (3[0])), and by four non-unique synapomorphies (e.g., pronotum with shallow longitudinal impression on disc (7[1]) and the penis with the subtriangular apex (35[0])).

Figures

FIGURE 1. Map of SouthAmerica with detailed area indicating the distributional records forPilielmis Hinton, 1971 (Coleoptera: Elmidae).

FIGURE 3. Pilielmis murcia Hinton, 1971 (Coleoptera: Elmidae). Adult male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 5. Pilielmis murcia Hinton, 1971 (Coleoptera: Elmidae). Adult, male. (A) Genitalia, ventral view. (B) Sternite VIII, dorsal view, with the anterior region highlighted. (C). Segment IX, ventral view. (D) Tergite VIII, dorsal view.

FIGURE 7. Pilielmis sara Hinton, 1971 (Coleoptera: Elmidae). Adult, male, habitus (Paratype). (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5mm.

FIGURE 9. Pilielmis sara Hinton, 1971 (Coleoptera: Elmidae). Adult, male. Paratype. (A) Genitalia, ventral view. (B) Tergite VIII, dorsal view. (C). Sternite VIII, dorsal view. (D) Metathoracic wing. (E) Segment IX, ventral view.

FIGURE 11. Pilielmis halia Hinton, 1971 (Coleoptera: Elmidae). Adult, male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 13. Pilielmis halia Hinton, 1971 (Coleoptera: Elmidae). Adult, male. (A) Genitalia, ventral view. (B) Tergite VIII, dorsal view. (C) Sternite VIII, dorsal view. (D) Metathoracic wing. (E) Segment IX, ventral view.

FIGURE 15. Pilielmis apama Hinton, 1971 (Coleoptera: Elmidae).Adult, male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 17. Pilielmis apama Hinton, 1971 (Coleoptera: Elmidae). Adult, male. (A) Genitalia, ventral view. (B) Sternite VIII, dorsal view. (C) Tergite VIII, dorsal view. (D) Metathoracic wing. (E) Segment IX, ventral view.

FIGURE 19. Pilielmis clita Hinton, 1971 (Coleoptera: Elmidae). Adult, male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 21. Pilielmis clita Hinton, 1971 (Coleoptera: Elmidae). Adult, male. (A) Genitalia, ventral view. (B) Sternite VIII, dorsal view. (C) Tergite VIII, dorsal view. (D) Segment IX, dorsal view. (E) Metathoracic wing.

FIGURE 23. Pilielmis abdera Hinton, 1971 (Coleoptera: Elmidae).Adult, male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 25. Pilielmis abdera Hinton, 1971 (Coleoptera: Elmidae). Adult, male. (A) Genitalia, ventral view. (B) Tergite VIII, dorsal view, with the latero-central region highlighted. (C) Segment IX, ventral view. (D) Sternum VIII, dorsal view. (E) Metathoracic wing.

FIGURE 27. Pilielmis shepardi Polizei & Barclay, 2019 (Coleoptera: Elmidae). Adult male, habitus. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 29. Pilielmis shepardi Polizei & Barclay, 2019 (Coleoptera: Elmidae). Adult male. (A) Genitalia, ventral view. (B) Sternite VIII fused with ventrite V, dorsal view. (C) Tergite VIII, dorsal view. (D). Segment IX, ventral view.

FIGURE 31. Pilielmis baniwa sp. nov. (Coleoptera: Elmidae). Adult male, habitus. Holotype. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5mm.

FIGURE 33. Pilielmis baniwa sp. nov. (Coleoptera: Elmidae). Adult male. Holotype. (A) Male genitalia, ventral view. (B) Sternite VIII, dorsal view. (C) Tergite VIII, dorsal view. (D) Metathoracic wing. (E) Segment IX, ventral view.

FIGURE 34. Pilielmis muricula sp. nov. (Coleoptera: Elmidae).Adult female. Holotype. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 37. Pilielmis nheengatu sp. nov. (Coleoptera: Elmidae). Adult male, habitus. Holotype. (A) Dorsal view. (B) Ventral view. (C) Lateral view. (D) Front view. Scale bars = 0.5 mm.

FIGURE 38. Pilielmis nheengatu sp. nov. (Coleoptera: Elmidae). Adult male, mouthparts. Paratype. (A) Labrum, dorsal view. (B) Maxilla, dorsal view. (C) Mandible, dorsal view. (D) Mandible ventral view. (E) Labium, ventral view. Scale bars = 0.1 mm.

FIGURE 39. Pilielmis nheengatu sp. nov. (Coleoptera: Elmidae). Adult male. Paratype. (A) Genitalia, ventral view. (B) Segment IX, ventral view. (C) Tergite VIII, dorsal view. (D) Sternum VIII, dorsal view. (E) Metathoracic wing.

FIGURE 40. Pilielmis nheengatu sp. nov. (Coleoptera: Elmidae). Adult female. Paratype. (A) Ovipositor, ventral view. (B) Tergite VIII, dorsal view. (C) Sternite VIII, dorsal view, with anterolateral region highlighted.

FIGURE 41. Paratypes of Pilielmis (Coleoptera: Elmidae). (A) Pilielmis murcia. (B) Pilielmis sara. (C) Pilielmis halia. (D) Pilielmis apama. (E) Pilielmis clita. (F) Pilielmis abdera. (G) Pilielmis shepardi. (A–F) Material housed at the NHMUK. (G) Material housed at MZSP. Scale bars = 0.5 mm.

FIGURE 42. Cladogram resulting from parsimony analysis under implied weights (K = 1) showing phylogenetic hypotheses for Pilielmis (Coleoptera: Elmidae) and related taxa. Black circles indicate unique apomorphies, white circles indicate non-unique changes on the tree, either forward or reverse. Numbers above and below circles are character and state numbers, respectively (see Appendix I). Letters (in green) above branches are node indication. Italicized numbers (in red) below branches indicate Relative Bremer values.

FIGURE 43. Strict consensus cladogram of the six most parsimonious trees resulting from the parsimony analysis under equal weights showing phylogenetic hypotheses for Pilielmis (Coleoptera: Elmidae) and related taxa.