Anchylorhynchus vittipennis Voss, 1943

De, Bruno A. S. & Vanin, Sergio A., 2020, Systematic revision and morphological phylogenetic analysis of Anchylorhynchus Schoenherr, 1836 (Coleoptera, Curculionidae: Derelomini), Zootaxa 4839 (1), pp. 1-98 : 62-91

Anchylorhynchus vittipennis Voss, 1943

( Figures 7D , 8Y , 14 B–C, 19E, 20F, 23F)

Ancylorrhynchus vittipennis Voss, 1943:64 (description in key). Holotype: Male ; São Paulo, Brazil; 1911; R. Fischer col. Deposited in DEI, examined.

Anchylorhynchus vittipennis ; Vaurie, 1954:38 (revision); Wibmer & O’Brien, 1986:196 (cat.).

Ancylorrhynchus nigripennis Voss, 1943:62 (description in key). Holotype destroyed ( Vaurie 1954).

Anchylorhynchys nigripennis ; Vaurie, 1954:38 (revision, synonymy to Anchylorhynchus vittipennis ); Wibmer & O’Brien, 1986:196 (cat.).

Ancylorrhynchus lineatus Bondar, 1943a:363 (description). Lectotype (designated by Vaurie, 1954): Male; Santa Catarina, Brazil; Maller col.; In “ Cocos romanzoffiana ”. Deposited in AMNH, examined.

Anchylorhynchus lineatus ; Blackwelder, 1947:827 (cat.); Vaurie, 1954:38 (revision, lectotype designated, synonymy to Anchylorhynchus vittipennis ); Wibmer & O’Brien, 1986:196 (cat.).

Redescription. Male: Body size (length of elytra + pronotum) 4.8–5.5 mm.

Head: Rostrum 1.4–1.6 times as long as pronotum; 1.2–1.3 times wider at apex than at base; integument brown to black; with seven longitudinal carinae (including a pair along scrobes), distinct throughout their length; scrobes parallel to rostrum; apex covered by microsetae; areas between dorsal carinae covered by wide yellowish scales, distinctively narrower towards the apex of rostrum, directed toward the central carina. Eyes 1.3–1.4 times as high as wide; 0.6–0.7 times more separated above than below. Head integument brown, similar to or lighter than rostrum; covered by large yellowish scales around the eyes and on the sides and dark microsetae elsewhere; scales directed to a point basal to the inter-ocular fovea. Antennae with scape straight, barely reaching anterior eye margin; funicle: antennomere I of funicle only slightly wider than II, II about as long as I, and about 1.5 times as long as III, VI longer than wide and narrower than club; club about as long as antennomeres IV–VI of the funicle. Left mandible slightly sinuate at outer margin; with one dorsal seta; outer tooth forming a sharp angle with the mandible margin, with somewhat blunt apex; inner tooth absent or weakly developed; molar region straight. Left maxilla with stipes strongly elongate, with a long ventral seta; galeo-lacinial complex not reaching the apex of palpomere I; palpiger with transverse ventral region, dorsal region with an acute angle at base; palpomere I with two apical setae shorter than the length of palpomere I; palpomere II approximately as long as palpomere I. Labium prementum distinctively narrower than postmentum, about 1.5 times as wide as long, lateral margins divergent in median region, with a right angle in lateral-apical region, with two oblique rows of setae in dorsal region; ligula shorter than half of the length of prementum; palps separated by about half the width of palpomere I; palpomere I transverse; palpomere II shorter than palpomere III; palpomere III with one lateral seta.

Thorax: Pronotum width at base 1.6–1.7 times pronotum length; base not lobed at middle; lateral margins in dorsal view convergent and regularly curved from base to apex, sometimes with a subtle apical constriction; integument brown, similar to head; covered by rows of yellow and black scales on sides and by yellowish microsetae on disc; scales on the disk directed to the center or to the apex. Profemur approximately 3 times as long as wide; dorsal margin with curvature about as pronounced as ventral margin; tooth well-defined; about 2.5 times as wide as protibia. Protarsus with tarsomere I about as long as tarsomere III; tarsomeres I and II longer than wide; with short and dense setae on the ventral surface, without sparse and long setae on the sides. Scutellum 0.6–0.8 times longer than wide; integument brown, lighter than elytra; scales brown. Elytra 1.5–1.6 times wider than pronotum; 1.4–1.5 times longer than wide; 3.6–4.0 times longer than pronotum; humeri rounded; lateral margins distinctively wider at middle; with uniformly black scales or with black scales on odd intervals and yellowish scales on even interval; epipleura with a distinct inflexion on interval IX, covered by dark scales. Ventral region of thorax integument yellowish to black; scales yellowish, similar yellowish scales in dorsal region. Hypomeron covered by sparse scales with truncated apex similar in shape to those in pronotum; scales next to coxal cavities larger. Prosternum length 0.9–1.2 times the width of coxae; postocular lobes indistinct, with apical-lateral margin of prothorax regularly curved to slightly sinuate; densely covered by mostly non-overlapping scales with truncated apex narrower than those in pronotum. Metepisternum covered by non-overlapping scales, or overlapping only next to the anterior margin. Metasternum central concavity distinct; with long setae in the center.

Abdomen: Ventrites III–V covered by erect setae. Aedeagus about 3 times as long as wide; about 5 times as long as high; slightly wider at opening; with a truncated anterior process; ventral plate thin and barely sclerotized, extending to approximately 0.2 times the length of apodemes. Apodemes about 3 times as long as aedeagus. Endophallus with one pair of membranous pockets containing microtrichae next to the basal sclerite; microtrichae of the apical region not organized in bands.

Female: Body size 4.9–5.6 mm. Rostrum 1.4–1.5 times longer than pronotum; width at apex 1.1–1.3 times width at base. Eyes 1.3–1.5 times as high as wide; 0.6–0.9 times more separated above than below. Pronotum 1.6–1.7 times as wide at base as long. Prosternum length 1.0–1.1 times the coxal width. Scutellum 0.8–1.0 times as long as wide. Elytra 1.3–1.7 times as wide as pronotum; 1.4–1.6 times as long as wide; 4.2–4.3 times as long as pronotum. Ventrites III–V with recumbent setae. Ventrites III and IV with posterior retraction slightly sinuate and reaching the middle of the segment.

Remarks. This species can be recognized by the very small scales in the disk of the pronotum, revealing the punctures underneath ( Figure 7D ) and by its color pattern with longitudinal rows of light scales on the sides of the pronotum, elytra black or with bands of yellowish scales along the even intervals ( Figures 14 B–C). Some specimens of Anchylorhynchus aegrotus and females of Anchylorhynchus vanini may have similarly reduced scales in the pronotum, but in these cases A. vittipennis can be distinguished by always having at least some black scales in the elytra, and by conical pronotum, with lateral margins always converging from the base and never with an inflexion in the middle ( Figure 8Y ). In A. aegrotus and A. vanini , the pronotal margins are parallel at base in males and have a strong inflexion in the middle in females ( Figures 8A, 8W ).

Geographical Distribution. This species occurs in southern Brazil, between the states of São Paulo and Santa Catarina ( Figure 24 ).

Host Plants. Possibly Syagrus romanzoffiana (Cham.) Glassman.

Bondar (1943a) received specimens for the type series of A. lineatus from southern Brazil, from A. Maller, J. Moure and R. Lange, collected in flowers of a palm, “evidently Cocos romanzoffiana Chamiso ” (junior synonym of Syagrus romanzoffiana (Cham.) Glassman ). There is no other record of host plant and we did not find A. vittipennis when sampling from Syagrus romanzoffiana elsewhere. This species is rare in collections when compared to the two other species associated with this palm ( A. variabilis and A. aegrotus ). It is possible that it is indeed associated with S. romanzoffiana , but with smaller populations and/or more restricted geographical distribution.

Examined Specimens. Type Material. BRAZIL. São Paulo: São Paulo , R. Fischer, III/1905 (♂ holotype Anchylorhynchus vittipennis DEI) . Santa Catarina: No locality, A. Maller, XI/1943 (♂ lectotype Anchylorhynchus lineatus AMNH); II/1943 (6♀, 1♂ paralectotypes Anchylorhynchus lineatus AMNH); XI/1943 (1♀, 1♂ paralectotypes Anchylorhynchus lineatus MNRJ) .

Other Material. BRAZIL. Paraná: Curitiba, Dept. Zoo., 25/XI/1967 (1♂ DZUP) ; Morretes, IAPAR, C. I. I. F., 24–25/XI/1984 (1♀ DZUP) . Santa Catarina: Cauna [Porto União, Santa Cruz do Timbó ], A. Maller, II/1946 (1♀ AMNH); XII/1945 (5♀ AMNH); Itajaí, collector illegible, VII /1919 (1♀ MZSP) ; No locality, no collector, no date (3♂ AMNH) ; Nova Teutônia [Seara, Nova Teutônia], F. Plaumann, 06/I/1946 (3♀, 1♂ AMNH); 15/XII/1946 (2♀ AMNH); 18/II/1946 (1♀, 1♂ AMNH); 20/XII/1940 (1♀ AMNH); 20/XII/1945 (3♀, 5♂ AMNH); 25/XII/1945 (2♀, 6♂ AMNH); 27/I/1946 (4♀, 1♂ AMNH); X/1941 (1♀ MZSP); XII/1945 (1♂ MZSP) ; Rio Vermelho , A. Maller, I/1945 (4♀, 2♂ AMNH) ; Dirings, II/1949 (1♀ MZSP); II/1953 (1♀, 1♂ MZSP); III /1957 (3♀, 3♂ MZSP) . São Paulo: Barueri, K. Lenko, 04/I/1967 (1♂ MZSP); 15/X/1960 (2♂ MZSP); XI/1965 (1♀ MZSP) ; São Paulo, Santana, F. Halik, 15/XII/1957 (1♂ USNM); 25/I/1962 (1♀ USNM) . NO COUNTRY. No region : No locality, Muller, no date (1♀, 1♂ CEAH) ; no collector, no date (1♂ MNRJ, 1♀ MZSP) .

Key to the species of Anchylorhynchus

1. Rostrum with 2–3 dorsal longitudinal carinae ( Figures 2 G–J).................................................. 2

- Rostrum with 7 dorsal longitudinal carinae ( Figures 2 A–F).................................................... 3

2. Length of elytra + length of pronotum greater than 5 mm; rostrum with 3 well-delimited carinae; scales of the disk of the pronotum directed to the apex ( Figure 8V ); sixth antennomere of the funicle longer than wide...... A. tricarinatus ( Figure 12E )

- Length of elytra + length of pronotum less than 5 mm; rostrum with 2–3 delimited carinae, sometimes weakly defined; scales of the disk of the pronotum directed to the center or the base ( Figure 8D ); antennomere VI of the funicle as long as wide.................................................................................. A. bicarinatus ( Figure 10G )

3. Scales along the longitudinal median line of the pronotum directed to the base, at least in the basal half (e. g. Figures 8C, 8F, 8P )................................................................................................. 4

- Scales along the longitudinal median line of the pronotum with a different configuration (e. g. Figures 8A, 8B, 8E )....... 12

4. Scape reaching or almost reaching the anterior margin of the eye (e. g. Figures 2D, 2H )............................. 5

- Scape clearly not reaching the anterior margin of the eye (e. g. Figure 2B )....................................... 10

5. Scape reaching well beyond the anterior margin of the eye ( Figures 2D , 12B ); carinae in the rostrum poorly defined near base ( Figures 2 C–D); eyes round; rostrum black and remainder of the body light yellow; males with rostrum, antennae and prolegs at least as long as the elytra; females with rostrum about as wide at the apex as at the base.... A. pinocchio ( Figures 12 B–C)

- Scape only reaching the the anterior margin of the eye ( Figure 2H ); carinae in the rostrum well delimited throughout ( Figures 2 A–B); eyes oval; color pattern variable................................................................... 6

6. Lateral margins of the pronotum converging at base ( Figures 8G, 8P ); pronotum transverse, at least 1.6 times wider at base than long................................................................................................ 7

- Lateral margins of the pronotum parallel at base, converging from the median region (e. g. Figures 8F, 8N, 8H ); pronotum more elongate, not more than 1.5 times as wide at base as long...................................................... 8

7. Large, length of elytra + length of the pronotum greater than 7 mm; profemur with a well-defined tooth ( Figure 18A ); scales of the basal half of the pronotum directed to the base ( Figure 8G ); hypomeron and mesepimeron with black scales ( Figures 11 A–B, 16B)................................................................. A. burmeisteri ( Figures 11 A–B)

- Length of elytra + length of the pronotum less than 7 mm; profemur with an indistinct tooth obscured by scales ( Figure 18C ); scales of most of the pronotum directed to the base ( Figure 8P ); ventral region without black scales ( Figure 12A )...................................................................................... A. multisquamis ( Figure 12A )

8. Rostrum shorter than the pronotum; second antennomere of the funicle shorter than the first; epipleura and a central band extending from the base of the pronotum to the base of the elytra with dark brown scales.............. A. bucki ( Figure 10L )

- Rostrum at least as long as the pronotum; second antennomere of the funicle as long as the first; uniform coloration or a different pattern........................................................................................... 9

9. Rostrum about 1.5 times the length of the pronotum; second antennomere of the funicle longer than the first; integument of head and rostrum dark brown to black, distinctly darker than the antennae and the rest of the body; scales with variable color patterns, yellow, brown or black................................................ A. luteobrunneus ( Figures 11 I–K)

- Rostrum only slightly longer than the pronotum; second antennomere of the funicle as long as the first; head and rostrum reddish to brown, with a similar color to the rest of the body; scales with uniform coloration, yellow to grayish.............................................................................................. A. camposi ( Figure 11C )

10. Elytra with lateral margins subparallel, not widened in the middle; protarsomere I distinctly longer than the third............................................................................................ A. rectus ( Figure 12E )

- Elytra with lateral margins at least slightly widened in the middle; protarsomere I about as long as the third............. 11

11. Eyes 0.6–0.9 times more separated above than below; pronotum with lateral margins slightly converging at base ( Figure 8C ); base of pronotum slightly lobed in the middle ( Figure 8C ); elytra usually widened in the middle; hypomeron sparsely covered by elongate scales............................................................. A. amazonicus ( Figures 10 D–F)

- Eyes 1.1–1.2 times more separated above than below; pronotum with lateral margins usually parallel at base ( Figure 8L ); base of pronotum almost straight ( Figure 8L ); elytra usually with subparallel margins; hypomeron densely covered by wide scales................................................................................. A. imitator ( Figure 11G )

12. Antennomeres I–III of the funicle approximately the same length and distinctly longer than the following ones ( Figures 3 F–G); integument sometimes dark, but scales always with uniform pale coloration...................................... 13

- Antennomeres I–III of the funicle about as long as the other antennomeres, or antennomeres III of the funicle distinctly shorter than the second; scales with variable coloration............................................................ 15

13. Lateral margins of the pronotum parallel at base ( Figure 8K ); prosternum covered by widely overlapping scales; pronotum less than 1.5 times as wide as long; rostrum not more than 1.6 longer than the pronotum................ A. goiano ( Figure 11F )

- Lateral margins of the pronotum converging at base ( Figures 8M, 8T ); prosternum covered by either sparse or overlapping scales; pronotum more than 1.5 times as wide as long at base; rostrum at least 1.6 times longer than the pronotum........ 14

14. Scape clearly not reaching the anterior margin of the eye (as in Figure 2B ); profemur, in lateral view, about 4 times as long as wide ( Figure 18J ); hypomeron with scales with similar size throughout; males with protarsomeres I–II extremely widened, dorsaly flattened and round ( Figure 18J ).................................................. A. latipes ( Figure 11H )

- Scape reaching or almost reaching the anterior margin of the eye (as in Figure 2H ); profemur, in lateral view, about 3 times as long as wide ( Figure 18H ); hypomeron with scales close to the coxal cavities larger; males with protarsomeres I–II cylindrical, only slightly wider than in females ( Figure 18H )...................................... A. trapezicollis ( Figure 12F )

15. Ventral region and epipleura covered with hairs ( Figure 16D )....................... A. variabilis ( Figures 13 H–L, 14A)

- Ventral region of thorax and epipleura covered with scales ( Figures 16 A–C)...................................... 16

16. Antennal club as long as the funicle ( Figure 3A )........................................ A. bicolor ( Figures 10 H–K)

- Antennal club shorter than the funicle.................................................................... 17

17. Scales of the disk of the pronotum clearly directed to the apex ( Figures 8A, 8O, 8Q ); body (except rostrum) covered by uniformly yellow or pale yellow scales; if a round patch of black scales present on the sides of the elytra (rare), then size minute, length of pronotum + elytra less than 4 mm ................................................................ 18

- Scales of the disk of the pronotum either directed to the center or base, or only a few scales directed to the apex ( Figures 8A, 8W ), or minute and barely visible ( Figures 7 C–D); body scales variable......................................... 20

18. Rostrum black with carinae poorly delimited near the base (as in Figures 2 C–D); areas between carinae covered with setae throughout the rostrum; pronotum with lateral margins converging at base ( Figure 8B )............ A. albidus ( Figure 10C )

- Rostrum brown, with similar color as the rest of the body, and with well-delimited carinae (as in Figures 2 A–B); areas between carinae covered with scales at base; pronotum with lateral margins parallel to subparallel at base ( Figures 8O, 8Q )....... 19

19. Length of pronotum + length of elytra less than 4 mm; first antennomere of the funicle distinctly wider than the second ( Figure 3D ); scales uniformly yellow, or rarely with a round patch of black scales on the side of each elytron; elytra more than 4 times longer than the pronotum............................................................ A. minimus ( Figure 11L )

- Length of pronotum + length of elytra greater than 5 mm; first antennomere of the funicle about as wide as the second; body reddish brown, uniformly colored and covered by pale yellow scales; elytra about 3 times as long as the pronotum........................................................................................... A. parcus ( Figure 12D )

20. Scales in the basal half of the pronotum clearly visible and directed to the center or the base ( Figure 8I ); lateral margins of the pronotum clearly converging at base and regularly curved to the apex ( Figure 8I ); male protarsi with elongate setae on the sides; color uniformly yellow........................................................ A. centrosquamatus ( Fig. 2.213 )

- Scales in the disk of the pronotum variably reduced ( Figures 7 C–D) or directed to the center or base ( Figures 8A, 8W, 8Y ); pronotum with variable shape; both sexes without long fringe of setae in protarsi; color variable …................... 21

21. Scales of the head sometimes very small and difficult to see. If visible, not directed to the interocular fovea, but to a point basal in relation to it (e. g. Figure 2E ); protarsomeres I–II longer than wide (as in Figure 18G ); epipleura usually with a distinct inflexion along interval IX.............................................................................. 22

- Scales of the head directed to the interocular fovea (as in Figures 2A, 2C, 2G ); protarsomeres I–II either as long as wide or wider than long (as in Figure 18A ); epipleura without a distinct inflexion........................................ 24

22. Rostrum at most 1.2 times longer than the pronotum; mandibles with a single long seta (as in Figure 4F ); integument yellow to light brown, never black; elytra with yellow scales or sometimes with a small patch of brown scales near the scutellum.................................................................................. A. aegrotus ( Figures 10 A–B)

- Rostrum longer than 1.2 times the pronotum; integument and scales with variable color: if integument uniformly yellow to light brown, then mandibles with two long setae each ( Figure 4H ).................................................. 23

23. Lateral margins of the pronotum regularly converging from the base ( Figure 8Y ); rostrum at least 1.4 times longer than the pronotum; integument black in rostrum, elytra and parts of legs and ventral region; elytra either entirely covered by black scales or with black scales intercalated by longitudinal bands of light scales in the even intervals..... A. vittipennis ( Figures 14 B–C)

- Lateral margins of the pronotum either parallel at base (males) or converging at base with a distinct inflexion in the middle (females) ( Figure 8W ); rostrum 1.2–1.4 times the length of the pronotum; males with uniformly yellow color, females yellow except for the pronotum disk with black integument and extremely reduced scales.............. A. vanini ( Figures 13 F–G)

24. Antennal club about as long as the last four antennomeres of the funicle ( Figure 3B ); prosternum and hypomeron with scales wider than those of the pronotum (as in Figure 9F ); rostrum at most 1.5 times as long as the pronotum; pronotum with a cross of black scales on the disk, surrounded by orange scales............................... A. chrysomeloides ( Figure 11E )

- Antennal club about as long as the last three antennomeres of the funicle; prosternum and hypomeron with more elongate scales than those of the pronotum, except near the coxae in males ( Figures 9 G–H); rostrum at least 1.5 times as long as the pronotum; color pattern variable, potentially with extreme polymorphism in a single population..................................................................................................... A. tremolerasi ( Figures 12 G–L, 13A–D)

Phylogeny

CHARACTERS

We were able to score 113 morphological characters in total for the 25 species of Anchylorhynchus and five outgroups. Eleven of these were discretized based on measurements, with the remainder being coded directly as discrete. These eleven discretized characters were the result of applying the filters described in the methods section to 38 possible measurements (table available https://github.com/brunoasm/revision_anchylorhynchus) and removing correlated characters or those that could be explained by a single Normal distribution. Fifty characters were binary, with 10 of them being autapomorphies, while 63 were multistate, with 25 of them being ordered. Missing or inapplicable data occupied 4.3% of the 3420 cells in the matrix, while polymorphic states occupied 4.04%. The annotated character matrices are available on Morphobank at http://morphobank.org/permalink/?P2797. Consistency index (CI) and retention index (CI) calculated on the parsimony topology. Discussions on reconstructions consider only parsimony-based reconstructions onto both tree topologies (i. e. parsimony and Bayesian).

Discrete characters

1. Rostrum, integument, color (unordered, CI 94, RI 33).

(0) yellowish.

(1) reddish ( Figure 2E ).

(2) reddish-brown ( Figure 2G ).

(3) black ( Figure 2F ).

This character was coded as polymorphic for most species. Exceptions are A. albidus and A. pinocchio , always with black rostrum. These species are sister groups in the parsimony topology, but not in the Bayesian topology.

2. Rostrum length, sexual dimorphism (binary, CI 100, RI 0).

(0) Males and females about the same size.

(1) Male rostrum much longer than female ( Figure 12 B–C).

Autapomorphy of Anchylorhynchus pinocchio .

3. Male, rostrum, longitudinal carinae, number (ordered, CI 71, RI 75).

(0) none.

(1) two ( Figure 2J ).

(2) three ( Figure 2G ).

(3) five.

(4) seven ( Figure 2A ).

Seven longitudinal carinae in males is a synapomorphy of Anchylorhynchus , with A. bicarinatus and A. tricatinatus forming a clade in which this number was reduced. Species of Celetes and Elaeidobius included in the analysis have five carinae, which is convergently evolved in some groups in Derelomini (character 47 in Franz 2006).

4. Male, rostrum, longitudinal carinae, whether distinct (unordered, CI 75, RI 50).

(0) all carinae distinct from base to apex ( Figure 2B ).

(1) two lateralmost carinae indistinct near base ( Figure 2D ).

(2) all carinae indistinct.

Carinae are distinct in most species of Anchylorhynchus and outgroups. The common ancestor of A. pinocchio , A. albidus and A. luteobrunneus shows a transition to indistinct carinae (states 1 and 2), with the former two sharing all carinae indistinct. This is a synapomorphy for the group under the parsimony topology. Celetes langei independently evolved indistinct carinae and some individuals of A. vanini show lateralmost carinae indistinct.

5. Rostrum, longitudinal carinae, sexual dimorphism (unordered, CI 100, RI 100).

(0) no dimorphism ( Figure 2 ).

(1) female with no carinae at base.

(2) female with no carinae throughout.

The lack of sexual dimorphism in rostral carinae is an unreversed synapomorphy of Anchylorhynchus .

6. Rostrum, scrobe, angle (binary, CI 50, RI 80).

(0) parallel to rostrum.

(1) directed obliquely downwards ( Figure 2B ).

The oblique scrobe is a synapomorphy of Anchylorhynchus , reversed in A. pinocchio . This condition is also found in other genera of Derelomini not sampled here (character 58, Franz 2006).

7. Male, rostrum, apex, vestiture, shape of scales (unordered, CI 67, RI 0).

(0) microsetae ( Figure 2A ).

(1) narrow elongate scales ( Figure 2I ).

(2) wide scales ( Figure 2G ).

Most species show microsetae between rostral carinae, but these transition to larger scales in the common ancestor of A. bicarinatus (state 1) and A. tricarinatus (state 2), and also independently in A. luteobrunneus (state 1). 8. Male, rostrum, dorsal scales, color (unordered, CI 85, RI 0).

(0) yellowish ( Figure 2G )

(1) orange.

(2) reddish-brown.

(3) black.

Yellowish represent the ancestral condition, with multiple independent origins of the other colors, none of them being a synapomorphy of smaller groups.

9. Male, rostrum, dorsal scales at the base, shape (unordered, CI 78, RI 60).

(0) narrow to setiform.

(1) wide scales ( Figure 2 ).

(2) microsetae.

Wide scales at the base of the rostrum is a synapomorphy of Anchylorhynchus , reversed to setiform scales independently in several species. State 2 is autapomorphic of Celetes langei .

10. Male, rostrum, dorsal scales, orientation (unordered, CI 100, RI 100).

(0) directed towards central carina ( Figure 2A ).

(1) directed towards base of rostrum ( Figure 2 G–J).

(2) not oriented.

Dorsal scales directed to the base of the rostrum are a synapomorphy of A. bicarinatus and A. tricarinatus . Unoriented scales are only found in Andranthobius bondari , which also lack dorsal carinae.

11. Male, rostrum, antennal insertions, position (binary, CI 50, RI 67).

(0) median.

(1) distal ( Figure 2 ).

Distal antennal insertions in males are shared between Anchylorhynchus and Celetes binotatus , being a synapomorphy of the group in the Bayesian topology but homoplastic in the parsimony topology. The rostrum with subapical antennal insertions (character 57 in Franz 2006) was recovered as a synapomorphy of Anchylorhynchus . This might only be a difference in coding of this character, since in Anchylorhynchus the insertions are even more apical than in Celetes binotatus . However, Valente (2005) found that the position of insertions varies among species of Celetes , being apical in many of them.

12. Head, integument, color difference to the rostrum (unordered, CI 91, RI 50).

(0) color similar to rostrum ( Figure 2E ).

(1) distinctly lighter than the rostrum ( Figure 2D ).

(2) distinctly darker than the rostrum.

The head and rostrum with similar color as ancestral for Anchylorhynchus , Celetes and Elaeidobius . There are multiple independent transitions to the other states in Anchylorhynchus , with the head distinctly lighter than the rostrum being a synapomorphy of A. pinocchio and A. albidus in the parsimony topology.

13. Head, dorsal region, vestiture, shape of scales (unordered, CI 75, RI 83).

(0) microsetae.

(1) with microsetae and large scales ( Figure 2E ).

(2) mostly covered by large scales ( Figure 2C ).

Head covered by at least some scales (states 1 and 2) are a synapomorphy of Anchylorhynchus and Celetes . Within Anchylorhynchus , the head entirely covered by scales in the ancestral condition, with scales on the sides and microsetae in the middle being a synapomorphy for a group formed by A. variabilis , A. vittipennis , A. aegrotus , A. vanini and A. bucki . In the latter this condition is polymorphic, some individuals having the head entirely covered by scales.

14. Head, dorsal region, vestiture, orientation of scales (unordered, CI 67, RI 85).

(0) directed to a point slightly more proximal than the interocular fovea ( Figure 2E ).

(1) directed to the base the rostrum ( Figure 2F )

(2) directed to the interocular fovea ( Figure 2A ).

The scales in the head directed to the interocular fovea is a synapomorphy of Anchylorhynchus unrecognized in previous studies. This condition is reversed to scales directed to a point more basal than the fovea in the taxon formed by A. variabilis , A. vittipennis , A. aegrotus , A. vanini and A. bucki . Scales directed to the base of the rostrum are an autapomorphy of A. burmeisteri . Franz (2006) found the presence of the fovea itself to be variable within Anchylorhynchus , but we found it to be present in all species.

15. Head, dorsal region, vestiture, color (unordered, CI 70, RI 40).

(0) uniformly yellowish ( Figure 2A ).

(1) uniformly dark.

(2) varying from dark to yellowish or orange.

The head covered by yellow scales or setae represents the ancestral condition. There are several transitions to darker colors, with this being a synapomorphy for A. variabilis , A. vittipennis and A. aegrotus in the parsimony topology, and this group plus A. vanini in the Bayesian topology.

16. Male, antennae, scape, length (unordered, CI 40, RI 63).

(0) not reaching the eye margin ( Figure 2B ).

(1) reaching the eye margin ( Figure 2H ).

(2) extending beyond the eye margin ( Figure 2D ).

The male scape reaching the eye margin or extending the eye margin is a synapomorphy of Anchylorhynchus + Celetes binotatus in the Bayesian topology, with this being a homoplasy in the parsimony tree. It is reversed in the monophyletic group formed by A. latipes , A. centrosquamatus , A. rectus , A. imitator , A. amazonicus and A. camposi , with the latter showing yet another reversal to state 1. The scape reaching beyond the eye margin is a synapomorphy of A. pinocchio .

17. Male, antennae, scape, shape (binary, CI 100, RI 0).

(0) straight ( Figure 3 A–D,F–I).

(1) curved ( Figure 3E ).

A distinctively curved scape is an autapomorphy of A. pinocchio .

18. Antennae, funicle, number of antennomeres (binary, CI 100, RI 100).

(0) 7.

(1) 6 ( Figure 3 ).

The funicle with 6 antennomeres is a synapomorphy of Anchylorhynchus . Within Derelomini , this is also found in Derelominus ( Franz 2006) , a genus not sampled here.

19. Left mandible, external margin, shape (unordered, CI 29, RI 44).

(0) not sinuate ( Figure 4A ).

(1) slightly sinuate ( Figure 4D ).

(2) strongly sinuate ( Figure 4E ).

The external margin not sinuate is the ancestral condition. A slightly sinuate margin is a synapomorphy of A. variabilis and A. vittipennis , but convergently present in several other species. A transition to strongly sinuate margin is found in the main clade of species associated with Butia palms: A. bicolor , A. chrysomeloides , A. burmeisteri , A. parcus , A. tremolerasi and A. multisquamis , with the latter representing a reversal to non-sinuate external margin of the mandible.

20. Left mandible, dorsal region, setae, number and shape (unordered, CI 100, RI 100).

(0) one very short seta.

(1) none.

(2) one long seta ( Figure 4A ).

(3) one long and one short seta ( Figure 4H ).

(4) three long setae ( Figure 4C ).

The mandible with a single long seta is a synapomorphy of Anchylorhynchus , with autapomorphic transitions to three long setae ( A. tricarinatus ) and one long and one short seta ( A. vanini ). A glabrous mandible was shared by the two species of Elaeidobius .

21. Left mandible, external tooth, orientation in relation to external margin (binary, CI 33, RI 0.5).

(0) evenly curved with the lateral margin ( Figure 4C ).

(1) sharply angled in relation to the lateral margin ( Figure 4E ).

In both trees, the ancestral condition is the tooth evenly curved with the lateral margin, with a sharply angled tooth appearing at least 5 times independently in both trees.

22. Left mandible, external tooth, shape (binary, CI 0.25, RI 0.4).

(0) pointed ( Figure 4C )

(1) rounded to truncated ( Figure 4D ).

The ancestral condition is a pointed external tooth. In both trees, a rounded or truncated tooth evolved independently in A. minimus , A. rectus and the common ancestor of A. vanini , A. bucki , A. aegrotus , A. variabilis and A. vittipennis , with the pointed tooth in the latter representing a reversion.

23. Left mandible, internal tooth, degree of development (binary, CI 100, RI 100).

(0) well developed ( Figure 4A ).

(1) indistinct to poorly developed ( Figure 4F ).

The ancestral condition in Anchylorhynchus is a mandible with two teeth, as found in most Derelomini ( Franz 2006) . In the parsimony tree, a reduction of the internal tooth is a synapomorphy of the clade formed by A. variabilis , A. vittipennis , A. aegrotus and A. bucki . In the Bayesian tree, A. vanini is nested within this clade and represents a reversion. Franz (2006) remarked that the internal tooth of A. variabilis is variably reduced (character 3 in Franz 2006).

24. Left mandible, molar region, shape (binary, CI 25, RI 50).

(0) straight ( Figure 4B ).

(1) sinuate ( Figure 4C ).

A straight molar region represents the ancestral condition, with four independent transitions to sinuate. One of them represents a synapomorphy for the clade A. amazonicus , A. imitator , A. camposi , A. rectus .

25. Left maxilla, stipes, shape (ordered, CI 33, RI 56).

(0) subquadrate ( Figure 5C ).

(1) moderately elongate ( Figure 5A )

(2) strongly elongate ( Figure 5I ).

Moderately elongate stipes are the ancestral condition for Anchylorhynchus , with several transitions to more elongate and less elongate. Subquadrate stipes are a synapomorphy of A. bicarinatus and A. tricarinatus , and also found in A. centrosquamatus . Strongly elongate stipes are a synapomorphy of A. variabilis + A. vittipennis and independently also of A. bicolor , A. chrysomeloides , A. burmeisteri , A. multisquamis , A. parcus and A. tremolerasi .

26. Left maxilla, stipes, ventral seta (binary, CI 50, RI 0).

(0) absent ( Figure 5A )

(1) present ( Figure 5B ).

A ventral seta is present is most Derelomini ( Franz 2006) . Here we found no seta in Elaeidobius subvittatus and Andranthobius bondari . Within Anchylorhynchus , the seta was absent in A. imitator and variable in its sister group A. amazonicus .

27. Left maxilla, galeo-lacinial complex, length in relation to maxillary palp (ordered, CI 17, RI 38).

(0) not reaching the apex of palpomere I ( Figure 5H )

(1) reaching the apex of palpomere I ( Figure 5I )

(2) reaching beyond the apex of palpomere I ( Figure 5A ).

This is a highly homoplastic character, with several transitions between states.

28. Left maxilla, palpiger, setae, arrangement (unordered, CI 100, RI 100).

(0) seven long and several short setae ( Valente & de Medeiros, 2013: Figure 2C ).

(1) only one seta.

(2) one long seta and several short setae ( Figure 5 ).

(3) two short setae of approximatelly equal length.

(4) three short setae of approximatelly equal length.

(5) two long and two short setae.

The palpiger with one long and several short setae is a synapomorphy of Anchylorhynchus , with a change to seven long setae in A. vanini . Franz (2006) also recovered this as a synapomorphy of the genus, convergent with Neopsilorhinus .

29. Left maxilla, palpiger, ventral region, shape (ordered, CI 40, RI 83).

(0) transverse ( Figure 5H ).

(1) subquadrate ( Figure 5C ).

(2) elongate ( Figure 5A ).

A subquadrate palpiger is found in Celetes and inferred to be the ancestral state for Anchylorhynchus . The first split within Anchylorhynchus marks divergent evolution of this character. The clade formed by species exclusively associated with Syagrus , including Anchylorhynchus amazonicus , A. albidus and other 9 species has an elongate palpiger as synapomorphy, reversed in A. camposi . In its sister group, containing all other species of Anchylorhynchus , there are two independent transitions to a transverse palpiger.

30. Left maxilla, palpiger, dorsal region, shape of proximal margin (unordered, CI 100, RI 100)

(0) evenly curved.

(1) with an obtuse angle ( Figure 5B ).

(2) with an acute angle ( Figure 5C ).

An evenly curved proximal margin of the palpiger was only observed in Andranthobius bondari . The ancestral condition in Anchylorhynchus , shared with the rest of the outgroup, is an obtuse angle, while a large clade including A. variabilis , and A. bicolor and 12 other species as an acute angle as a synapomorphy.

31. Left maxilla, palpomere I, apical setae, number (unordered, CI 60, RI 83).

(0) none ( Figure 5D ).

(1) one.

(2) two ( Figure 5C )

(3) three ( Figure 5B ).

The palpomere I with two or three setae is a synapomorphy for Anchylorhynchus , also found in Franz (2006), but with a transition to glabrous in A. minimus . Two setae is the inferred ancestral condition for the genus, with three setae being a synapomorphy for the clade A. amazonicus , A. imitator , A. camposi , A. rectus , A. centrosquamatus , A. latipes , A. trapezicollis and A. goiano , and also convergently found in A. vanini .

32. Left maxilla, palpomere I, apical setae, length (binary, CI 14, RI 40).

(0) elongate ( Figure 5A ).

(1) short ( Figure 5E ).

This is a highly homoplastic character, both within Anchylorhynchus and in the outgroup. The first split within Anchylorhynchus separates one clade mostly with elongate setae (including A. amazonicus and A. albidus ) and a clade mostly with short setae ( A. tremolerasi and A. aegrotus ).

33. Left maxilla, palpomere I, shape (ordered, CI 40, RI 72).

(0) transverse.

(1) about as wide as long ( Figure 5G ).

(2) longer than wide ( Figure 5A ).

The outgroup has clearly transverse palpomeres, with elongate palpomeres being a synapomorphy of Anchylorhynchus . There are a few transitions to less elongate palmpomeres, but never as wide as in other genera.

34. Left maxilla, palpomere II, length in relation to palpomere I (ordered, CI 28, RI 44).

(0) longer than palpomere I ( Figure 5A ).

(1) about as long as palpomere I ( Figure 5B ).

(2) shorter than palpomere I ( Figure 5C ).

Highly homoplastic character, with several transitions between states 0 and 1. A palpomere II shorter than the palmpomere I is an autapomorphy of A. tricarinatus . Characters 27 and 28 in Franz (2006) deal with relations between palpomere II and palpomere I of the maxilla, but are not directly equivalent to this character. He reports that a palpomere II distinctly slender and more elongate than the palpomere I is a synapomorphy of Anchylorhynchus , which is consistent with our findings with the exception of A. tricarinatus .

35. Labium, prementum, aspect ratio (ordered, CI 38, RI 58).

(0) about 2 times as wide as long ( Figure 6B ).

(1) about 1.5 as wide as long ( Figure 6A ).

(2) about as long as wide ( Figure 6D ).

(3) longer than wide.

With exception of A. pinocchio , Anchylorhynchus shares with Celetes langei and Andranthobius bondari a clearly transverse prementum, with the aspect ratio varying within the genus. In A. pinocchio , the prementum is longer than wide.

36. Labium, prementum, lateral margins, shape (binary, CI 25, RI 25).

(0) diverging in the median region ( Figure 6B ).

(1) parallel in the median region ( Figure 6A ).

The lateral margins of the prementum parallel are inferred to be the ancestral condition. Diverging margins are a synapomorphy for A. variabilis + A. vittipennis , convergently found in two other species of Anchylorhynchus and also in Andranthobius bondari .

37. Labium, prementum, apical-lateral region, shape (unordered, CI 22, RI 53).

(0) retracted ( Figure 6F ).

(1) forming a right angle ( Figure 6I ).

(2) lobed ( Figure 6A ).

This is a highly homoplastic character, with all three states found in Anchylorhynchus .

38. Labium, prementum, central longitudinal band of setae (binary, CI 50, RI 85).

(0) absent ( Figure 6A ).

(1) present ( Figure 6F ).

The ancestral condition in Anchylorhynchus and in the outgroup is absent. There are two transitions inferred in Anchylorhynchus , with differences between the Bayesian and the parsimony tree. With the parsimony topology, the presence of a longitudinal band of setae in the prementum is a synapomorphy for one of the two clades diverging at the base of the genus, reversed in the common ancestor of A. variabilis , A. vittipennis , A. aegrotus and A. bucki . Under Bayesian inference, A. vanini is nested within the latter clade, rendering a convergence between A. vanini and the other species with these setae the preferred optimization.

39. Labium, prementum, lateral oblique band of setae (binary, CI 25, RI 75).

(0) absent ( Figure 6C ).

(1) present ( Figure 6A ).

The presence of these obliquely oriented setae is a synapomorphy for one of the large clades diverging at the base of the genus, including A. amazonicus and A. luteobrunneus . It is also convergently found in A. variabilis + A. vittipennis , A. tricarinatus and A. bicolor . Character 33 in Franz (2006) corresponds to these setae and is deemed a synapomorphy of Anchylorhynchus . It is possible that this character corresponds to both character 38 and 39 as delimited here, but even considering both together there is variation within the genus, with A. aegrotus and A. bucki not having either longitudinal or oblique rows of setae.

40. Labium, prementum, apical-lateral setae, number (unordered, CI 100, RI 100).

(0) one

(1) 2–4

(2) more than 4 ( Figure 6 ).

More than four apical-lateral setae in the prementum is a synapomorphy of Anchylorhynchus + Celetes .

41. Labium, prementum, ligula, degree of development (ordered, CI 50, RI 81).

(0) about as long as the prementum.

(1) about 0.5 times the length of prementum ( Figure 6D ).

(2) present but less than 0.5 times the length of prementum ( Figure 6J ).

(3) present and barely visible ( Figure 6F ).

(4) absent ( Figure 6G ).

The ligula (anterior tendon in Franz 2006) is about as long as the prementum in Elaeidobius and Andranthobius , with a shorted ligula being a synapomorphy of Celetes + Anchylorhynchus . Within Anchylorhynchus , there are several transitions in the degree of development, but it is never longer than 0.5 the length of the prementum.

42. Labium, prementum, distance between palpi (ordered, CI 22, RI 50).

(0) less than half the width of palpomere I ( Figure 6F ).

(1) about half the width of palpomere I ( Figure 6H ).

(2) about the same as the width of palpomere I ( Figure 6J ).

This is a highly homoplastic character, with the 3 states being found in Anchylorhynchus .

43. Labium, palpomere I, shape (ordered, CI 20, RI 43).

(0) wider than long ( Figure 6C ).

(1) about as wide as long ( Figure 6D ).

(2) longer than wide ( Figure 6A ).

This is a highly homoplastic character, with the 3 states being found in Anchylorhynchus and the outgroup.

44. Labium, palpomere I, proximal ventral setae, number (unordered, CI 67, RI 75).

(0) one long seta

(1) three long setae ( Valente & de Medeiros, 2013: Figure 2D ).

(2) one long seta and many shorter ones ( Figure 6 ).

The presence of at least three setae (regardless of length) is a synapomorphy of Anchylorhynchus . Most species of Anchylorhynchus present state 2 (also found in character 41 in Franz 2006), with state 1 being an autapomorphy of A. vanini convergently found in Celetes langei .

45. Labium, palpomere II, length in relation to palpomere III (ordered, CI 25, RI 25).

(0) shorter than palpomere III ( Figure 6I ).

(1) about as long as palpomere III ( Figure 6A ).

(2) longer than palpomere III ( Figure 6F ).

This is a highly homoplastic character, not defining many groupings.

46. Labium, palpomere III, lateral setae, number (unordered, CI 33, RI 69).

(0) none ( Figure 6F )

(1) one ( Figure 6C )

(2) two ( Figure 6A ).

Franz (2006) in character 45 found that the presence of one lateral seta in palpomere III was convergently present in a clade including Anchylorhynchus and Celetes and in other Derelomini . Here we additionally find that Celetes langei and Anchylorhynchus tremolerasi present transitions to no seta, and that a large clade including A. amazonicus and A. goiano have two setae, a condition convergently evolved in A. tricarinatus .

47. Pronotum, proximal margin, shape (unordered, CI 33, RI 33).

(0) not lobed in the middle ( Figure 8S ).

(1) slightly lobed in the middle ( Figure 8T ).

(2) strongly lobed in the middle ( Figure 8V ).

This is a highly homoplastic character both in Anchylorhynchus and in the outgroup.

48. Male, pronotum, lateral margins, curvature (binary, CI 33, RI 43).

(0) evenly curved ( Figure 8A ).

(1) with an inflexion in median region ( Figure 8J ).

A pronotum with an inflexion in the middle is convergently found in four different groups within Anchylorhynchus under the parsimony topology. Under the Bayesian topology, this is a synapomorphy of A. pinocchio + A. luteobrunneus , with a convergent transition in a clade including A. tricarinatus , A. bicarinatus , A. multisquamis , A. burmeisteri , A. parcus , A. tremolerasi , A. chrysomeloides and A. bicolor , with a reversion in the latter.

49. Male, pronotum, lateral margins, direction at base (unordered, CI 38, RI 47).

(0) diverging.

(1) parallel ( Figure 8A ).

(2) subparallel ( Figure 8L ).

(3) converging ( Figure 8B ).

This is a highly homoplastic character with multiple transitions within Anchylorhynchus . A round pronotum with margins diverging at base is found only in Celetes .

50. Pronotum, lateral margins, curvature, sexual dimorphism (unordered, CI 67, RI 0).

(0) No dimorphism

(1) Males evenly curved and females with an inflexion in median region.

(2) Males with an inflexion in median region and females evenly curved.

State 2 is an autapomorphy of Anchylorhynchus luteobrunneus and state 1 is shared between A. vanini and A. aegrotus , being a synapomorphy in the Bayesian topology and convergent in the parsimony topology.

51. Pronotum, lateral margins, direction at base, sexual dimorphism (binary, CI 20, RI 0).

(0) No dimorphism.

(1) Females with margins more convergent than males.

Under the parsimony topology, there are 5 convergent origins of this sexual dimorphism. Under the Bayesian topology, there are 4, with one of them being a synapomorphy of A. aegrotus and A. vanini .

52. Male, pronotum, apical constriction (binary, CI 73, RI 57).

(0) with a slight constriction ( Figure 8B )

(1) without constriction ( Figure 8A ).

Seven species of Anchylorhyncus were polymorphic for this character, with several convergent origins of the pronotal constriction in Anchylorhynchus .

53. Male, pronotum, setae, shape (unordered, CI 83, RI 50).

(0) with scales on the sides and microsetae on the disc ( Figure 7 C–D)

(1) scales ( Figure 7A )

(2) long setae

(3) short setae and narrow scales

(4) short setae.

The pronotum covered with scales is a synapomorphy of Anchylorhynchus , with a modification to disk covered by microsetae in A. vittipennis and A. aegrotus (polymorphic).

54. Pronotum, vestiture, sexual dimorphism (binary, CI 100, RI 0).

(0) not sexually dimorphic.

(1) males with scales and females with microsetae on the disk.

The sexually dimorphic pronotum vestiture is an autapomorphy of A. vanini .

55. Pronotum, setae, color, sexual dimorphism (binary, CI 100, RI 0).

(0) not sexually dimorphic (e. g. Figures 13 H–L).

(1) sexually dimorphic, with males always yellow ( Figures 13 F–G).

The sexually dimorphic pronotum color is an autapomorphy of A. vanini .

56. Pronotum, setae along the longitudinal midline (unordered, CI 20, RI 47).

(0) directed towards the base ( Figure 8C ).

(1) directed towards the apex ( Figure 8B ).

(2) directed towards the center ( Figure 8A ).

This is a highly homoplastic character with several transitions within Anchylorhynchus . While Vaurie (1954) identified that the variation is useful to diagnose species, it does not form clear groupings for phylogenetics.

57. Male, profemora, lateral view, shape (unordered, CI 1, RI 0).

(0) about equally curved in dorsal and ventral margins ( Figure 18B ).

(1) distinctly more straight on dorsal margin ( Figure 18A ).

(2) distinctly more straight on ventral margin.

State 1 is an autapomorphy of A. burmeisteri and State 2 is an autapomorphy of C. binotatus .

58. Male, profemora, ventral tooth, degree of development (unordered, CI 40, RI 40).

(0) absent.

(1) poorly developed ( Figure 18C ).

(2) well developed ( Figure 18A ).

The presence of a ventral tooth in the profemora is a synapomorphy of Celetes + Anchylorhynchus . There are multiple transitions to a poorly developed tooth within Anchylorhynchus , but it is always observed. Franz (2006), character 90, observed that femoral teeth are convergently present in other genera of Derelomini as well.

59. Male, protarsi, tarsomere I, length in relation to tarsomere III (ordered, CI 40, RI 63).

(0) approximately as long as tarsomere III ( Figure 18C ).

(1) slightly longer than tarsomere III ( Figure 18I )

(2) more than 3 times longer than tarsomere III ( Figure 18E ).

The tarsomere I slightly longer than the tarsomere III is the ancestral condition in Anchylorhynchus , shared with Celetes and Andranthobius . There are three convergent transitions to tarsomeres I and III of approximately the same length, in the most recent common ancestor of A. bicolor , A. bucki and 12 other species, in the clade of A. amazonicus , A. imitator and A. camposi , and independently in A. centrosquamatus . A tarsomere I much longer than the tarsomere III is an autapomorphy of A. pinocchio .

60. Male, protarsi, tarsomeres I and II, shape (ordered, CI 36, RI 53).

(0) distinctly longer than wide ( Figure 18E ).

(1) slightly longer than wide ( Figure 18F ).

(2) as long as wide ( Figure 18B ).

(3) slightly wider than long ( Figure 18C ).

(4) distinctly wider than long ( Figure 18J ).

While useful to diagnose species, this character is highly homoplastic and presents multiple transitions within the genus.

61. Female, protarsi, tarsomeres I and II, shape (ordered, CI 22, RI 50).

(0) longer than wide ( Figure 18I )

(1) as long as wide ( Figure 18B )

(2) slightly wider than long ( Figure 18C ).

Because the shape of tarsi in females seems to be somewhat uncorrelated from females, we coded this as a separate character. As is the case for males, it resulted to be highly homoplastics with multiple transitions between states.

62. Male, protarsi, fringe of ventral long setae (binary, CI 33, RI 80).

(0) absent ( Figure 18A ).

(1) present ( Figure 18B ).

Males in some species have a fringe of lateral long setae in the protarsi. The presence of these setae is a synapomorphy of one of the major clades in Anchylorhynchus , including A. amazonicus and A. luteobrunneus . The only exception is A. albidus , which is inferred as a reversal in the parsimony topology, but not on the Bayesian topology, in which A. albidus is sister to this group. The long setae are also convergently found in A. parcus .

63. Scutellum, vestiture, color (unordered, CI 78, RI 0).

(0) uniformly yellow to orange ( Figure 11D ).

(1) bicolor ( Figure 11A ).

(2) uniformly dark ( Figure 11E ).

The ancestral state is the uniformly yellow to orange scutellum. Bicolor or uniformly dark evolved independently several times in Anchylorhynchus .

64. Elytra, humeri shape (binary, CI 13, RI 50).

(0) with a marked angle ( Figure 12E )

(1) rounded ( Figure 12F ).

The humeri always present a marked angle in the outgroup, with the ancestral state for Anchylorhynchus being ambiguous and several transitions within the genus. Somewhat rounded humeri are part of the rounded body aspect considered as a synapomorphy of Anchylorhynchus (character 1 in Franz 2006).

65. Elytra, lateral margins, shape (binary, CI 25, RI 50).

(0) wider in the middle ( Figure 12F ).

(1) subparallel ( Figure 12E ).

Celetes and Elaeidobius have subparallel elytral margins, and within Anchylorhynchus there are several transitions to a more rounded profile, with the elytra wider in the middle. This is part of the rounded body aspect considered as a synapomorphy of Anchylorhynchus (character 1 in Franz 2006).

66. Elytra, dorsal region, vestiture, color (unordered, CI 93, RI 50).

(0) uniformly yellow to orange ( Figure 13E )

(1) bicolor ( Figure 13B )

(2) uniformly dark ( Figure 13C ).

There are at least 5 independent transitions to bicolor or dark elytra in Anchylorhynchus , but in most of these cases the species is polymorphic also having uniformly yellow individuals, which inflates consistency index. Most of these transitions are concentrated in the clade including A. variabilis and A. bicolor .

67. Elytra, dorsal region, vestiture, whether sexually dimorphic (binary, CI 100, RI 0).

(0) not sexually dimorphic

(1) sexually dimorphic, with males always yellow ( Figure 13 H–L).

Sexually dimorphic coloration in the elytra is an autapomorphy of A. variabilis .

68. Elytra, epipleura, vestiture, color (binary, CI 60, RI 50).

(0) similar to the disc ( Figure 13G )

(1) with a stripe of dark scales ( Figure 13H ).

There are several independent transitions dark epipleura in Anchylorhynchus , and in some cases the trait is polymorphic. Most of these transitions are concentrated in the clade including A. variabilis and A. bicolor .

69. Elytra, epipleura, degree of inflexion (binary, CI 75, RI 50).

(0) with a smooth inflexion ( Figure 10D ).

(1) with a sharp inflexion ( Figure 10A ).

A sharp inflexion in the epipleura has convergently evolved in A. trapezicollis and in the clade of A. variabilis , A. vittipennis , A. aegrotus , A. vanini and A. bucki . The latter represents a reversal to rounded inflexion in the parsimony topology or the ancestral condition in the Bayesian topology.

70. Ventral region of the thorax, vestiture, color in relation to dorsal region (binary, CI 27, RI 38).

(0) similar ( Figure 10C ).

(1) distinctly lighter-colored ( Figure 10D ).

This is a highly homoplastic character with several transitions within Anchylorhynchus and in the outgroup.

71. Hypomeron, vestiture, shape of scales (unordered, CI 63, RI 75).

(0) microsetae.

(1) setiform ( Figure 15D ).

(2) acuminate ( Figure 9C ).

(3) truncated ( Figure 9B ).

The presence of acuminate or truncated scales in the hypomeron is a synapomorphy of Anchylorhynchus , with a reversal to setae in A. variabilis . There are multiple transitions between acuminate and truncated scales within the genus.

72. Hypomeron, region further from coxal cavities, vestiture, shape and size in of scales in relation to those in pronotum (unordered, CI 30, RI 13).

(0) distinctly more elongate.

(1) slightly more elongate ( Figure 9G ).

(2) similar ( Figure 9E ).

(3) wider ( Figure 9F ).

Inapplicable in the outgroup, since it lacks scales in the pronotum. This character is useful to diagnose species, but presents multiple transitions within Anchylorhynchus .

73. Male, hypomeron, region next to coxal cavities, vestiture, shape and size of scales (binary, CI 14, RI 57).

(0) similar ( Figure 9F ).

(1) wider ( Figure 9H ).

As character 72, this character is useful to diagnose species, but it is highly homoplastic and presents multiple transitions within Anchylorhynchus .

74. Hypomeron, region next to coxal cavities, vestiture, sexual dimorphism (binary, CI 100, RI 0).

(0) not sexually dimorphic.

(1) scales more elongate in females than in males ( Figures 9 G–H).

This is an autapomorphy of A. tremolerasi .

75. Prosternum, postocular lobes (binary, CI 100, RI 0).

(0) absent ( Figure 9A ).

(1) present ( Figure 9C ).

This is a synapomorphy of A. bicarinatus and A. tricarinatus .

76. Prosternum, elongate apical setae (binary, CI 100, RI 100).

(0) absent.

(1) present ( Figure 15 ).

The elongate setae on the sides and ventral side of the prothorax are a synapomorphy of Anchylorhynchus , also recognized by Franz (2006) in character 73 in his matrix. It is also present in some species of Celetes .

77. Prosternum, vestiture, density (unordered, CI 22, RI 50).

(0) very sparse.

(1) non-overlapping to slightly overlapping ( Figure 15B ).

(2) widely overlapping ( Figure 15A ).

A dense vestiture in the prosternum is found in C. binotatus and in Anchylorhynchus , which is a convergence in the parsimony topology and a synapomorphy in the Bayesian topology. There are multiple transitions between states 1 and 2 within Anchylorhynchus .

78. Prosternum, vestiture, shape and size of scales in relation to those in pronotum (unordered, CI 50, RI 50).

more elongate (0).

similar (1).

wider (2).

Inapplicable in the outgroup, since it lacks scales in the pronotum. The ancestral state is inferred as more elongate scales in the prosternum, with wider scales in the prosternum being a synapomorphy of A. bicarinatus and A. tricarinatus . Scales similar in size in the prosternum and pronotum are a synapomorphy of A. bicolor , A. chrysomeloides , A. multisquamis , A. parcus and A. burmeisteri , with a reversal in the latter. In the Bayesian topology, A. tremolerasi is nested in this group and also represents a reversal. It also evolved independently in 3 other species: A. imitator , A. centrosquamatus and A. luteobrunneus .

79. Prosternum, scales, apex, shape (unordered, CI 67, RI 83).

(0) microsetae

(1) setiform ( Figure 15D ).

(2) acuminate ( Figure 15A )

(3) truncated ( Figure 15C ).

The presence of scales in the prosternum is a synapomorphy of Anchylorhynchus . Large setae convergently evolved in A. variabilis and Celetes binotatus . The ancestral state for the genus is inferred to be acuminate scales, with truncated scales convergently evolving in at three to five groups, depending on the tree topology considered. In both topologies, it is an apomorphy of the monophyletic group including A. bicolor , A. chrysomeloides , A. burmeisteri , A. multisquamis , A. parcus and A. tremolerasi .

80. Prosternum, distance between coxal cavities (binary, CI 100, RI 100).

(0) separate.

(1) contiguous ( Figure 15 ).

Contiguous coxae are a synapomorphy of Anchylorhynchus and Celetes . The broader sample of Derelomini in character 81 in Franz (2006) reveals that this is actually the ancestral condition in the tribe, with transitions to separated coxae in several lineages.

81. Metepisternum, vestiture, density (binary, CI 11, RI 20).

(0) non-overlapping ( Figure 16A ).

(1) overlapping ( Figure 16B ).

This is a useful diagnostic character, with overlapping scales or setae in the metepisternum evolving multiple times within Anchylorhynchus and Celetes .

82. Male, metasternum, concavity, sharpness of edges (unordered, CI 25, RI 40).

(0) indistinct.

(1) with smooth edges.

(2) with sharp edges ( Figure 17 A–C).

A distinct metasternal concavity in males is the apomorphic state for Anchylorhynchus and Celetes . Sharp edges in this concavity is the inferred ancestral state for Anchylorhynchus , with two independent transitions to in- distinct concavity and four to a concavity with smooth edges.

83. Male, metasternum, concavity, vestiture, shape of setae (unordered, CI 20, RI 43).

(0) microsetae ( Figure 17C ) (1) long setae ( Figure 17 A–B).

(2) scales.

The vestiture in male metasternal concavity is a good diagnostic character, but highly homoplastic with several transitions within the genus and the outgroup.

84. Female, metasternum, vestiture, shape of setae (unordered, CI 17, RI 33).

(0) microsetae.

(1) long setae ( Figure 17D ).

(2) scales.

As is the case for males, in females the vestiture of the metasternum is highly homoplastic.

85. Male, ventrites III–V, vestiture, arrangement and shape of setae (unordered, CI 38, RI 47).

(0) recumbent setae ( Figure 19A ).

(1) erect hairs ( Figure 19E )

(2) elongate scales ( Figure 19B )

(3) elongate scales and recumbent hairs.

(4) microsetae.

As is the case for the vestiture in the metasternum, this character is highly homoplastic and presents several transitions.

86. Female, ventrites III–V, vestiture, arrangement and shape of setae (unordered, CI 60, RI 82).

(0) recumbent setae.

(1) elongate scales.

(2) microsetae.

The vestiture in female ventrites is less homoplastic when compared to males. In the Bayesian topology, there are two convergent transitions to elongate scales, in the monophyletic group including A. amazonicus and A. latipes , and in the group including A. bicolor , A. tremolerasi , A. tricarinatus and A. bicarinatus . In the parsimony topology, A. bicarinatus and A. tricarinatus are not nested in the latter group, representing a third transition. State 2 was only found in Andranthobius bondari .

87. Female, ventrites III–IV, degree of retraction (unordered, CI 33, RI 70).

(0) not retracted.

(1) slightly retracted ( Figure 20A ).

(2) strongly retracted ( Figure 20E ).

The ventrites III–IV in females with at least some degree of retraction are a synapomorphy of Anchylorhynchus . There are multiple transitions within the genus between states 1 and 2. It is suggested that this trait might be involved in palm pollen transfer (de Medeiros et al. 2019), and pollen is often found in the retraction in pinned specimens. This retraction was coded as character 114 in Franz (2006), also found as a synapomorphy of Anchylorhynchus .

88. Female, ventrites III–IV, shape of posterior margin (unordered, CI 50, RI 0).

(0) non-sinuate ( Figure 20B ).

(1) slightly sinuate ( Figure 20F ).

(2) strongly sinuate ( Figure 20G ).

A sinuate retraction convergently evolved in A. tremolerasi , A. burmeisteri . The state of the common ancestor of A. vittipennis + A. variabilis is ambiguous, with state 1 in the former and 2 in the latter. Franz (2006) considered the ventral female retraction in Anchylorhynchus to be always sinuate (character 114), but more sampling in the genus reveals that this is the least common state.

89. Male genitalia, aedeagus, aperture, shape (unordered, CI 33, RI 33).

(0) narrower than the anterior region

(1) as wide as the anterior region ( Figure 21A ).

(2) slightly wider than the anterior region ( Figure 21B ).

(3) strongly wider than the anterior region ( Figure 23E ).

There are multiple transitions between states 1 and 2 within Anchylorhynchus , with state 3 being an autapomorphy of A. variabilis .

90. Male genitalia, aedeagus, shape of apex (unordered, CI 40, RI 50).

(0) not constricted ( Figure 23B ).

(1) constricted and acute, round or truncated ( Figure 23A ).

(2) constricted and bilobed ( Figure 23E ).

Within Anchylorhynchus , a constricted apex in the aedeagus is the ancestral state. In the parsimony topology, this constriction was lost in the common ancestor of A. bicolor and A. tremolerasi , with state 1 in A. burmeisteri being a reversal. A bilobed apex is an autapomorphy of A. variabilis .

91. Male genitalia, aedeagus, ventral sclerotization, size (binary, CI 100, RI 100).

(0) not extending beyond the insertion of apodemes.

(1) extending beyond the insertion of apodemes ( Figures 21–23 ).

The presence of a ventral sclerotization visibly extending beyond the insertion of the apodemes is a synapomorphy of Anchylorhynchus and Celetes . This character was not scored by Franz (2006).

92. Male genitalia, aedeagus, ventral plate, degree of sclerotization (binary, CI 20, RI 60).

(0) well sclerotized ( Figure 21G )

(1) poorly sclerotized ( Figure 21A ).

Inapplicable in Elaeidobius and Andranthobius . Within Anchylorhynchus , there are at least 4 transitions to a ventral region in the aedeagus poorly sclerotized.

93. Male genitalia, endophallus, lateral pockets with microtrichiae (binary, CI 17, RI 55).

(0) absent ( Figure 21A ).

(1) present ( Figure 21E ).

The lateral regions with microtichiae are only found in Anchylorhynchus , but there are multiple transitions between states in the genus.

94. Male genitalia, endophallus, proximal complex sclerite (binary, CI 100, RI 100).

(0) absent.

(1) present ( Figures 21–23 ).

This proximal complex sclerite is a synapomorphy of Anchylorhynchus , also found by Franz (2006) in character 138.

95. Male genitalia, endophallus, paired distal sclerites (binary, CI 50, RI 88).

(0) absent ( Figure 22E ).

(1) present ( Figure 22 B–C).

Some species present a pair of sclerites distal and clearly separated from the proximal sclerite found in all species of Anchylorhynchus . This is a synapomorphy of the monophyletic group including A. amazonicus and A. goiano , convergently evolved in A. multisquamis .

96. Male genitalia, endophallus, paired distal sclerites, shape (unordered, CI 100, RI 100).

(0) L-shaped ( Figure 22C ).

(1) rounded ( Figure 22B )

(2) elongate ( Figure 22G ).

Inapplicable with species with state 0 in character 95. A simple elongate sclerite is an autapomorphy of A. multisquamis . A rounded sclerite evolved in the most recent common ancestor of A. amazonicus and A. goiano , with a further transition to an L-shaped sclerite being a synapomorphy of A. amazonicus , A. imitator , A. camposi , A. rectus and A. centrosquamatus .

97. Male genitalia, endophallus, hook-shaped sclerite (binary, CI 100, RI 0).

(0) absent ( Figure 21–23 ).

(1) present.

This sclerite is an autapomorphy of Elaeidobius kamerunicus .

98. Male genitalia, endophallus, round-shaped distal sclerite (binary, CI 100, RI 0).

(0) absent ( Figure 21–23 ).

(1) present.

This sclerite is an autapomorphy of Celetes langei .

99. Male genitalia, endophallus, distal sclerite shaped like a double hook (binary, CI 100, RI 0).

(0) absent ( Figure 21–23 ).

(1) present.

This sclerite is an autapomorphy of Andranthobius bondari .

100. Male genitalia, endophallus, distal slightly sclerotized area (binary, CI 100, RI 0).

(0) absent ( Figure 21E ).

(1) present ( Figure 21D ).

This is an autapomorphy of A. bicarinatus .

101. Male genitalia, endophallus, microtrichiae (binary, CI 100, RI 100).

(0) present ( Figures 21–23 )

(1) absent.

The absence of microtrichiae is a synapomorphy of Elaeidobius .

102. Male genitalia, endophallus, microtrichiae, arrangement (unorderd, CI 80, RI 67).

(0) all within the aedeagus

(1) not organized in stripes ( Figure 21B ).

(2) organized in a pair of longitudinal bands in the apical region ( Figure 21E ).

(3) organized in a pair of longitudinal bands in the median region ( Figure 21A ).

(4) organized in one transverse band in the apical region ( Figure 23C ).

In the outgroup, microtrichiae, when present are always in a distal position, close to the aperture and clearly within the limits of the aedeagus. In Anchylorhynchus , the microtrichiae in the endophallus extend beyond the limits of the aedeagus (states 2–4). These microtrichiae are ancestrally not organized in stripes, with state 3 being an autapomorphy of A. aegrotus and state 4 an autapomorphy of A. tricarinatus . State 2 is a synapomorpy of A. bicolor , A. chrysomeloides and A. tremolerasi in the Bayesian topology, and convergently present in the latter in the parsimony topology.

Discretized characters

103. Body size (geometric mean of pronotum length, pronotum width, elytra length, elytrum width) (ordered, CI 94, RI 0).

(0) body size measurement is about 1.04.

(1) body size measurement is about 1.86.

(2) body size measurement is about 2.75.

Most species have intermediate size (state 1), with multiple species in Anchylorhynchus being polymorphic between states 1 and 2. State 0 is only found in the outgroup.

104. Male, eyes above in relation to head width (unordered, CI 65, RI 14).

(0) eyes above measurement is about 0.18 times head width.

(1) eyes above measurement is about 0.26 times head width.

(2) eyes above measurement is about 0.32 times head width.

State 1 is a synapomorphy of Anchylorhynchus , but with multiple transitions within the genus.

105. Male, rostrum width base in relation to rostrum width apex (binary, CI 100, RI 100).

(0) rostrum width base measurement is about 0.76 times rostrum width apex.

(1) rostrum width base measurement is about 0.97 times rostrum width apex.

The male rostrum clearly wider at apex is a synapomorphy of Anchylorhynchus , with A. pinocchio being polymorphic.

106. Female, rostrum length in relation to the body size (geometric mean of pronotum length, pronotum width, elytra length, elytrum width) (ordered, CI 88, RI 0).

(0) rostrum length measurement is about 0.61 times the body size.

(1) rostrum length measurement is about 0.91 times the body size.

(2) rostrum length measurement is about 1.50 times the body size.

A long rostrum in females (state 2) is only found in the outgroup. State 1 is the ancestral for Anchylorhynchus , with convergent transitions to an even shorter rostrum (state 0) in A. aegrotus and A. bicolor .

107. Male, pronotum length in relation to the body size (geometric mean of pronotum length, pronotum width, elytra length, elytrum width) (binary, CI 100, RI 100).

(0) pronotum length measurement is about 0.61 times the body size.

(1) pronotum length measurement is about 0.79 times the body size.

A short pronotum (state 0) is a synapomorphy of Anchylorhynchus , with some species being polymorphic. This character has not been scored for other Derelomini in Franz (2006) .

108. Male, scutellum length in relation to the body size (geometric mean of pronotum length, pronotum width, elytra length) (binary, CI 100, RI 100).

(0) scutellum length measurement is about 0.05 times the body size.

(1) scutellum length measurement is about 0.09 times the body size.

A relatively long scutellum (state 1) is a synapomorphy of Anchylorhynchus . This character has not been scored for other Derelomini in Franz (2006) .

109. Male, scutellum width in relation to the body size (geometric mean of pronotum length, pronotum width, elytra length). (binary, CI 57, RI 40)

(0) scutellum width measurement is about 0.08 times the body size.

(1) scutellum width measurement is about 0.12 times the body size.

The outgroup has relatively narrow scutellum (state 0), with multiple transitions between the two states within Anchylorhynchus .

110. Male, eye height in relation to head width (binary, CI 57, RI 40).

(0) eye height measurement is about 0.45 times head width.

(1) eye height measurement is about 0.61 times head width.

A relatively narrow eye (state 1) is found in all species of Anchylorhynchus , but also in one species of Celetes and one of Elaeidobius .

111. Male, elytra length in relation to the body size (geometric mean of pronotum length, pronotum width, elytra length, elytrum width) (binary, CI 80, RI 0).

(0) elytra length measurement is about 1.85 times the body size.

(1) elytra length measurement is about 2.22 times the body size.

Species of Anchylorhynchus and Celetes have relatively long elytra (state 1), with polymorphism in some species of the former. State 0 is convergently found in Andranthobius bondari and Elaeidobius kamerunicus .

112. Male, prosternum length in relation to the pronotum size (geometric mean of pronotum length, pronotum width) (binary, CI 100, RI 100).

(0) prosternum length measurement is about 0.29 times the pronotum size.

(1) prosternum length measurement is about 0.55 times the pronotum size.

The short prosternum in relation to the pronotum is a synapomorphy of Anchylorhynchus . In Franz (2006), character 77, only the length of prosternum and mesosternum was compared, representing a different character.

113. Male, coxa width in relation to the pronotum size (geometric mean of pronotum length, pronotum width) (binary, CI 100, RI 0).

(0) coxa width measurement is about 0.19 times the pronotum size.

(1) coxa width measurement is about 0.29 times the pronotum size.

Small coxae (state 0) are an autapomorphy of Elaeidobius kamerunicus .

TREES

In the parsimony analysis, all values of k implying a mean distortion of at least 79.6 resulted in the same single most parsimonious tree. This corresponds to values of k of 6.3 and above, being the most stable region of the space of distortion. We therefore selected this tree, with an unweighted length of 661, consistency index of 52 and retention index of 57. Bayesian analysis inferred a 95% highest posterior density interval for α in the range of 1.35–3.76, indicating significant asymmetry in transition rates across characters. Both parsimony and Bayesian inference ( Figure 25 ) recover Anchylorhynchus as monophyletic with 100% support (bootstrap or posterior probability, respectively). There are 19 unreversed synapomorfies for Anchylorhynchus with unambiguous optimization ( Figure 26 ). Nine of these synapomorphies have not experienced additional transitions within the genus and are diagnostic for all species: (char. 5) lack of sexual dimorphism in number and distinctiveness of dorsolateral carinae in rostrum; (char. 18) antennal funicle with 6 antennomeres (also recovered in Franz, 2006); (char. 53) pronotum covered by scales at least on the sides; (char.77) prosternum with elongate apical setae (also recovered in Franz, 2006); (char. 87) ventrites III–IV of females with a retraction; (char. 95) male genitalia with complex apical sclerite (also recovered in Franz, 2006); (char. 107) short pronotum in males in relation to body size; (char. 109) wide scutellum in relation to body size; (char. 113) short prosternum in relation to body size. Most of the synapomorphies of Anchylorhynchus in Franz (2006) also resulted as synapomorphies of the genus here, even if some show additional transitions within the group. Exceptions are: (char. 39) the oblique bands of setae in the prementum, which were present in the species studied by Franz (2006), but exhibit multiple gains and losses in Anchylorhynchus and (char. 31) the number of apical-lateral setae in palpomere I, which is 2–3 in most species but reversed to none in A. minimus . We have not scored the character “body round”, but some of the species of Anchylorhynchus , such as A. rectus , are more elongate and could arguably be considered as elongate as some species of Phytotribus or Celetes . We have also not scored characters from the spiculum gastrale, since its morphology is invariable within the genus.

Under Bayesian inference, but not under parsimony, Celetes was found to be paraphyletic with respect to Anchylorhynchus , a result previously found in analyses focused on Celetes ( Franz & Valente 2005; Valente 2005). In a phylogeny of the whole Derelomini ( Franz 2006) , Celetes was recovered together with Phytotribus but not paraphyletic with respect to Anchylorhynchus . Most of the clades within Anchylorhynchus have low support in at least one of the trees, but there are several similarities between the parsimony and the Bayesian trees ( Figure 25 ). A notable difference is the position of the clade formed by A. tricarinatus and A. bicarinatus , which is well-supported regardless of the inference method. Another major difference is whether A. pinocchio , A. luteobrunneus and A. albidus constitute a monophyletic group (Bayesian tree) or not (parsimony tree). The common ancestor of Anchylorhynchus is inferred to have been associated with Syagrus , with this association being maintained in one of the two major clades in the genus ( Figure 27 ). In the other clade, there were at least two transitions to Butia , one transition each to Oenocarpus and Euterpe , and likely a transition back to Syagrus in Anchylorhynchus multisquamis , assuming that the genus host was correctly identified. Anchylorhynchus parcus , the only species with no host records, is nested within a group interacting with Butia .

Figures

FIGURE 2. Heads of species of Anchylorhynchus. Scale: 500 µm. A A. centrosquamatus, male, showing five well-defined dorsal carinae in the rostrum and scales on the head directed to the interocular fovea. B A. centrosquamatus, male, showing the pairs of lateral carinae along the scrobe, totaling 7 carinae in the rostrum. C A. pinocchio, female, showing five dorsal carinae poorly defined towards the base. D A. pinocchio, female, showing the lateral carinae poorly defined towards the base. E A. aegrotus, showing scales in the head smaller and directed to the base of the head. F A. burmeisteri, showing scales in the head directed to the rostrum. G A. tricarinatus, male, showing three well-defined dorsal carinae. H A. tricarinatus, male, showing no lateral carinae. I A. bicarinatus, male, with three well-defined dorsal carinae. H A. bicarinatus, male, with only two dorsal carinae and no central carina.

FIGURE 3. Antennae of males of species of Anchylorhynchus. Scales: 500 µm. A A. bicolor. B A. chrysomeloides. C A. variabilis D A. minimus. E A. pinocchio. F A. trapezicollis. G A. latipes. H A. rectus. I A. albidus.

FIGURE 4. Left mandibles of species of Anchylorhynchus (ventral view). Scales: 100 µm. A A. amazonicus. B A. trapezicollis. C A. tricarinatus D A. minimus. E A. tremolerasi. F A. variabilis. G A. aegrotus. H A. vanini.

FIGURE 5. Left maxillae of species of Anchylorhynchus (ventral view). Scales: 50 µm. A A. amazonicus. B A. trapezicollis. C A. tricarinatus. D A. minimus. E A. tremolerasi. F A. bucki. G A. aegrotus. H A. vittipennis. I A. variabilis.

FIGURE 6. Labia of species of Anchylorhynchus (dorsal view). Scales: 50 µm. Palp setae illustrated only on left side and lateral-apical setae of prementum only on right side. A A. amazonicus. B A. trapezicollis. C A. minimus. D A. pinocchio. E A. tricarinatus. F A. tremolerasi. G A. bicolor. H A. bucki. I A. aegrotus J A. variabilis.

FIGURE 7. Prothorax of species of Anchylorhynchus, dorsal view. Arrows show microsetae on disk. Scales: 500 μm. A A. trapezicollis. B A. bicarinatus. C A. aegrotus. D A. vittipennis.

FIGURE 8. Prothorax of all Anchylorhynchus species. Except for A. parcus, only known from the male holotype, the thorax of a male (left) and a female (right) are shown for each species. Arrows show the direction of scales in each position. Scale: 500 µm. A A. aegrotus. B A. albidus. C A. amazonicus. D A. bicarinatus. E A. bicolor. F A. bucki. G A. burmeisteri. H A. camposi. I A. centrosquamatus (from de Medeiros & Núñez-Avellaneda 2013). J A. chrysomeloides. K A. goiano. L A. imitator. M A. latipes. N A. luteobrunneus (from de Medeiros & Núñez-Avellaneda 2013). O A. minimus. P A. multisquamis. Q A. parcus. R A. pinocchio (from de Medeiros & Núñez-Avellaneda 2013). S A. rectus. T A. trapezicollis U A. tremolerasi V A. tricarinatus W A. vanini X A. variabilis Y A. vittipennis.

FIGURE 9. Prothorax of species of Anchylorhynchus, lateral view. Scales: 500 µm. A A. camposi. B A. rectus, showing larger scales next to coxal cavities. C A. bicarinatus, showing scales in the hypomeron larger than in the pronotum. D A. tricarinatus, showing scales in the hypomeron larger than in the pronotum. E A. goiano, showing larger scales next to coxal cavities. F A. multisquamis, showing dense vestiture in the hypomeron. G A. tremolerasi, female. H A. tremolerasi, male, showing sexual dimorphism with larger scales next to coxal cavities.

FIGURE 10. Habitus of species of Anchylorhynchus. Some images are mirrored in relation to originals. Scales: 1 mm. A A. aegrotus (male) B A. aegrotus (female) C A. albidus D A. amazonicus (male holotype) E–F A. amazonicus (Canaã dos Carajás, Brazil) G A. bicarinatus H–K A. bicolor. Photographs in K provided by Roberta Valente based on specimens loaned from NHMUK. L A. bucki.

FIGURE 11. Habitus of species of Anchylorhynchus (continued). Some images are mirrored in relation to originals. Scales: 1 mm. A–B A. burmeisteri. C A. camposi. D A. centrosquamatus (from de Medeiros & Núñez-Avellaneda 2013). E A. chrysomeloides. F A. goiano. G A. imitator. H A. latipes I–K A. luteobrunneus (from de Medeiros & Núñez-Avellaneda 2013). L A. minimus.

FIGURE 12. Habitus of species of Anchylorhynchus (continued). Some images are mirrored in relation to originals. Scales: 1 mm. A A. multisquamis. B A. pinocchio (male). C A. pinocchio (female). D A. parcus (male holotype). Photographed by Johannes Bergsten © 2017 Naturhistoriska riksmuseet. Background removed from original photo, light levels and contrast adjusted. Made available by the Swedish Museum of Natural History under CC-BY 4.0 international license. E A. rectus. F A. trapezicollis. G–L A. tremolerasi.

FIGURE 13. Habitus of species of Anchylorhynchus (continued). Scales: 1 mm. A–D A. tremolerasi. E A. tricarinatus. F A. vanini (male). G A. vanini (female). H A. variabilis (male). I–L A. variabilis (female).

FIGURE 14. Habitus of species of Anchylorhynchus (continued). Scales: 1 mm. A A. variabilis (female). B–C A. vittipennis.

FIGURE 15. Prosterna of species of Anchylorhynchus, ventral view. Scales: 500 µm. A A. bicarinatus. B A. camposi. C A. chrysomeloides. D A. variabilis.

FIGURE 16. Meso- and metathorax of species of Anchylorhynchus, lateral view. Scales: 500 µm. A A. camposi. B A. burmeisteri. C A. multisquamis. D A. variabilis.

FIGURE 17. Metathorax of species of Anchylorhynchus, ventral view. Scales: 500 µm. A A. trapezicollis. B A. aegrotus C A. vanini, male. D A. vanini, female.

FIGURE 18. Prolegs of species of Anchylorhynchus. Scales: 500 µm. A A. burmeisteri, male. B A. camposi, male. C A. multisquamis, male. D A. minimus, male. E A. pinocchio, male. F A. tricarinatus, male. G A. variabilis, male. H A. trapezicollis, male. I A. trapezicollis, female. J A. latipes, male. K A. latipes, female.

FIGURE 19. Male ventrites of species of Anchylorhynchus. Scales: 500 µm. A A. amazonicus, showing recumbent hairs. B A. amazonicus, showing a morph with less elongate scales. C A. imitator, showing that scales on ventrites are never as elongate as in A. amazonicus. D A. goiano. E A. vittipennis. F A. variabilis.

FIGURE 20. Ventrites of species of Anchylorhynchus. Retraction of ventrites III–IV is shown in grey. Scales: 500 µm. A A. amazonicus, female. B A. bucki, female. C A. bicarinatus, female. D A. bicarinatus, male. E A. bicolor, female. F A. vittipennis. G A. variabilis, female. H A. variabilis, male.

FIGURE 21. Male genitalia of species of Anchylorhynchus. A. parcus is known only from the holotype, which was not dissected. Scales: 500 µm. A A. aegrotus B A. albidus C A. amazonicus D A. bicarinatus E A. bicolor. F A. bucki. G A. burmeisteri. H A. camposi. I A. centrosquamatus (reproduced from de Medeiros & Núñez-Avellaneda, 2013).

FIGURE 22. Male genitalia of species of Anchylorhynchus (continued). Scales: 500 µm. A A. chrysomeloides B A. goiano C A. imitator D A. latipes E A. luteobrunneus (reproduced from de Medeiros & Núñez-Avellaneda, 2013). F A. minimus. G A. multisquamis. H A. pinocchio (reproduced from de Medeiros & Núñez-Avellaneda, 2013). I A. rectus.

FIGURE 23. Male genitalia of species of Anchylorhynchus (continued). Scales: 500 µm. A A. trapezicollis B A. tremolerasi C A. tricarinatus D A. vanini (digitally inked based on camera lucida illustration by Daniela Bená) E A. variabilis F A. vittipennis.

FIGURE 24. Geographical distribution of species of Anchylorhynchus based on specimens examined, excluding suspicious locations as indicated in the main text. The distribution of A. parcus is unknown. A A. aegrotus, A. albidus, A. amazonicus, A. centrosquamatus. B A. bicarinatus, A. bicolor, A. burmeisteri, A. rectus. C A. bucki, A. goiano, A. imitator, A. luteobrunneus. D A. camposi, A. minimus, A. multisquamis, A. tremolerasi. E A. latipes, A. pinocchio, A. trapezicollis, A. vittipennis. F A. chrysomeloides, A. tricarinatus, A. vanini, A. variabilis.

FIGURE 25. Phylogeny of Anchylorhynchus, including outgroups. A Bayesian tree with posterior probabilities as branch support. B Parsimony tree with symmetric bootstrapping as branch support.

FIGURE 26. Unambiguous character reconstructions on the parsimony tree.

FIGURE 27. Ancestral state reconstruction for the association with host plant genus on the Bayesian tree.