Anchylorhynchus tremolerasi Hustache, 1937

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 : 51-55

Anchylorhynchus tremolerasi Hustache, 1937

( Figures 4E , 5E , 6F , 8U , 9 G–H, 12G–L, 13A–D, 23B)

Ancylorrhynchus tremolerasi Hustache, 1937:9 (description); Bondar, 1943a:361 (revision). Holotype: Male; Prado, Montevideo, Uruguay; 31-XII-1908; collection A. Hustache; In flowers of “ Cocos pulposa B. R”. Deposited in MNHN, images examined.

Anchylorhynchus tremolerasi ; Blackwelder, 1947:827 (cat.); Vaurie, 1954:35 (revision); Viana, 1975:25, Fig. 1 (revision); Wibmer & O’Brien, 1986:196 (cat.)

Ancylorrhynchus pictipennis Hustache, 1937:10 (description); Bondar, 1943a:361 (revision) [new synonym]. Holotype: Male; Prado, Montevideo, Uruguay; 31-XII-1908; collection A. Hustache; In flowers of “ Cocos pulposa B. R”. Deposited in MNHN, images examined.

Anchylorhynchus pictipennis ; Blackwelder, 1947:827 (cat.); Vaurie, 1954:36 (revision); Viana, 1975:24 ( Fig. 14 ) (revision); Wibmer & O’Brien, 1986:196 (cat.)

Ancylorrhynchus hatschbachi Bondar, 1943a:363 (description). Lectotype (designated by Vaurie, 1954): Male; Curitiba, Paraná, Brazil; Bondar col.; In “ Cocos eriospatha ”. Deposited in AMNH, examined.

Anchylorhynchus hatschbachi ; Blackwelder, 1947:827 (cat.); Vaurie, 1954:32, Fig. 3G (revision, lectotype designation); Wibmer & O’Brien, 1986:196 (cat.); de Medeiros et al., 2014:18 (larval description, synonymy to Anchylorhynchus eriospathae )

Ancylorrhynchus erospathae Bondar, 1943a:364 (description, lapsus) [new synonym]. Lectotype (designated by Vaurie, 1954): Male; Curitiba, Paraná, Brazil; Bondar col.; In “ Cocos eriospatha ”. Deposited in AMNH, exammined.

Anchylorhynchus erospathae ; Blackwelder, 1947:827 (cat.); Wibmer & O’Brien, 1986:196 (cat.)

Anchylorhynchus eriospathae ; Blackwelder, 1947:827 (cat.); Vaurie, 1954:33 (revision, lectotype designated); Wibmer & O’Brien, 1986:196 (cat.); Franz, 2006:224 (phylogeny); de Medeiros et al., 2014:6 (larval description).

Redescription. Male: Body size (length of elytra + pronotum) 4.6–7.8 mm.

Head: Rostrum 1.5–1.8 times as long as pronotum; 1.1–1.3 times wider at apex than at base; with 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 yellowish to black scales, scales wide and distinctively narrower towards the apex of rostrum, directed toward the central carina. Eyes 1.4–1.7 times as high as wide; 0.7–1.0 times more separated above than below. Head with integument black to brown, similar to or darker than rostrum; entirely covered by black to yellow scales directed 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 as wide as long and narrower than club; club about as long as antennomeres IV–VI of funicle. Left mandible strongly sinuate at outer margin; with one dorsal seta; outer tooth forming a sharp angle with the mandible margin, with somewhat blunt apex; inner tooth well-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 slightly narrower than postmentum, about 2 times as wide as long, lateral margins parallel in median region, angled in lateral-apical region, with a median longitudinal row of short setae in dorsal region; ligula inconspicuous; palps separated by less than half the width of palpomere I; palpomere I transverse; palpomere II longer than palpomere III; palpomere III without lateral setae.

Thorax: Pronotum width at base 1.6–1.9 times pronotum length; base slightly lobed at middle; lateral margins in dorsal view subparallel at base and regularly curved to apex, sometimes with a subtle inflexion in the median region, with or without out a subtle apical constriction; integument black in areas with dark scales and brown otherwise; entirely covered by scales with diverse color patterns (see Figures 12 G–L, 13A–D); scales on disk directed 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 as long as wide; with short and dense setae on the ventral surface, without sparse and long setae on the sides. Scutellum 0.6–0.9 times longer than wide; integument brown to black; scales yellow or black. Elytra 1.2–1.6 times wider than pronotum; 1.4–1.9 times longer than wide; 3.5–4.2 times longer than pronotum; humeri rounded; lateral margins distinctively wider at middle; integument and scales with diverse color patterns (see Figures 12 G–L, 13A–D); epipleura without a marked inflexion on interval IX, with a band of dark scales or following the color pattern of disk. Ventral region of thorax integument yellowish to black; scales with variable colors, from white to black. Hypomeron covered by sparse scales, with most of the integument visible; covered by scales narrower than or similar to those in pronotum, with truncated apex; scales next to coxal cavities larger. Prosternum length 0.8–1.0 times the width of coxae; postocular lobes indistinct, with apical-lateral margin of prothorax regularly curved to slightly sinuate; covered by overlapping scales with truncated apex, narrower than those in pronotum. Metepisternum entirely covered by overlapping scales. Metasternum central concavity distinct; with very short setae in the center.

Abdomen: Ventrites III–V covered by narrow scales. Aedeagus about 3 times as long as wide; about 5 times as long as high; slightly wider at opening; without an anterior process; ventral plate strongly sclerotized, extending to approximately 0.4 times the length of apodemes. Apodemes about 1.5 times as long as aedeagus. Endophallus with two pairs of membronous pockets containing microtrichae next to basal sclerite; microtrichae of the apical region organized in a pair of bands, apparently the result of folds in membranes in this area.

Female: Body size 3.6–6.6 mm. Rostrum 1.5–1.7 times longer than pronotum; width at apex 1.0–1.3 times width at base. Eyes 1.3–1.7 times as high as wide; 0.8–1.0 times more separated above than below. Pronotum 1.6–1.9 times as wide at base as long. Proepisternum with scales narrower than those in pronotum. Prosternum length 0.7–1.0 times the coxal width. Scutellum 0.7–1.0 times as long as wide. Elytra 1.3–1.8 times as wide as pronotum; 1.4–1.7 times as long as wide; 3.7–4.9 times as long as pronotum. Ventrites III and IV with posterior retraction slightly sinuate and reaching the middle of the segment.

Remarks. This is the only species of Anchylorhynchus with sexual dimorphism in the size of scales of the proepisternum, with those of males being larger near the coxal cavities ( Figure 9 G–H). Some specimens can be distinguished by their unique color pattern ( Figures 12 G–L, 13A–D), and additionally this species can be recognized by a combination of characters: rostrum at least 1.5 times longer than the pronotum; scales of the disk of the pronotum directed to the center or apex ( Figure 8U ); mandible with external margin strongly sinuate ( Figure 4E ); antennal club about as long as the three last antennomeres of the funicle (similar to Figure 3B ).

Here we synonymize two formerly valid species with A. tremolerasi , after already having synonymized A. hatschbachi with A. eriospathae in a previous contribution ( de Medeiros et al. 2014). All of these species were previously distinguished by their color pattern alone. As previously accepted, A. eriospathae corresponds to Figures 12L and 13B, A . hatschbachi to Figures 12 I–J, A. pictipennis to Figure 12H, A . tremolerasi to Figure 12G and the other color patterns were undescribed. We could not find any other diagnostic character except for differences in average size, and many color morphs are often found together in a single inflorescence. Together, A. tremolerasi and A. pictipennis range between 4.7–7.8 mm in the length of pronotum + elytra, while A. eriospathae varies between 3.6–7.2 mm. While there is a difference in averages, there is also a large overlap. We dissected the male genitalia of each previously recognized species and did not find any differences. The sexual dimorphism in the size of prosternal scales close to the coxal cavities, larger in males, is unique to A. tremolerasi in the broad sense used here and shared by all populations in this species. There are multiple color morphs in a single population, including intermediate forms between the previously recognized species ( Figure 12K is a combination of A. hatschbachi , Figure 12J , and A. eriospathae , Figure 12L ). The type of A. pictipennis , for example, clearly has an integument color that follows the same pattern of scales of the type of A. tremolerasi . For these reasons, here we consider all of these color morphs sharing the diagnostic characters described in the previous paragraph to be the same species. Since A. tremolerasi and A. pictipennis were described together in the same article, there is no priority between both. Considering that not all specimens show the white-speckled scales in the elytra after which A. pictipennis is named, we consider A. tremolerasi to be a more appropriate accepted name.

Geographical Distribution. This species is distributed through Uruguay, eastern Argentina and southern Brazil, in coastal dunes, Araucaria forest and grasslands where Butia palms occur ( Figure 24 ). This species can also be found outside its native range, being dispersed together with their host palms, which are used for landscaping and transported as mature individuals extracted from their native habitats. Since A. tremolerasi pupates in the litter trapped by persistent leaf bases ( de Medeiros et al. 2014), weevils are transported together with the host plants.

Host Plants. Butia catarinensis Noblick & Lorenzi , Butia odorata (Barb. Rodr.) Noblick and Butia eriospatha (Mart. ex Drude) Becc.

Possibly a flower visitor of Phoenix canariensis Hort.

Hustache (1937) described A. tremolerasi and its junior synonym A. pictipennis based on material sampled from Cocos pulposa Barb. Rodr. , a synonym of Butia odorata (Barb. Rodr.) Noblick. Bondar described the junior synonyms A. eriospathae and A. hatschbachi from specimens collected from flowers of Cocos eriospatha Mart. , junior synonym of Butia eriospatha (Mart. ex Drude) Becc. We collected from Butia catarinesis Noblick & Lorenzi and from Butia eriospatha outside its native range, in the state of São Paulo, Brazil. Viana (1975) claims to have collected this species from flowers of Phoenix canariensis Hort , an introduced species in the New World.

Examined Specimens. Type Material. URUGUAY. Montevideo: Prado, J. Tremoleras, 31/XII/1908 (♂ holotype Anchylorhynchus tremolerasi MNHN, 1♀ paratype Anchylorhynchus tremolerasi MNHN, ♂ holotype Anchylorhynchus pictipennis MNHN, 1♀ paratype Anchylorhynchus pictipennis MNHN) .

BRAZIL. Paraná: Curitiba, G. Hatschbach, no date (♂ lectotype Anchylorhynchus eriospathae AMNH, 1♀ paralectotype Anchylorhynchus eriospathae CEAH, ♂ lectotype Anchylorhynchus hatschbachi AMNH, 6♀, 6♂ paralectotypes Anchylorhynchus hatschbachi AMNH, 2♀ paralectotypes Anchylorhynchus hatschbachi MLPA); B. Pohl, I/1943 (2♂ paralectotypes Anchylorhynchus hatschbachi MZSP) .

Note: the type series of A. hatschbachi have inconsistent labels, with specimens at the MZSP collected by B. Pohl and other labelled as collected by G. Hatschbach. Bondar (1943a) states that samples were sent to him by G. Hatschbach, but provides no information on the original collector.

Other Material. ARGENTINA. Buenos Aires: Buenos Aires, M. J. Viana, II/1950 (1 sex undetermined MLPA) ; no collector, 14/XII/1942 (1♂ AMNH) ; 30/XII/1942 (1♀ AMNH) ; Buenos Aires, San Fernando, Daguerre, XII/1957 (1 sex undetermined USNM) ; XII/1962 (1 sex undetermined USNM) .

BRAZIL. Bahia: No locality [mistake], no collector, no date (2♀ MNRJ) . Paraná: Curitiba, Casagrande, 17/XII/1980 (1♂ DZUP) ; 27/XII/1980 (11♀ DZUP) ; collector illegible, I/1944 (3♀, 1 sex undetermined MLPA) ; no collector, 01/I/1946 (1♀, 1♂ AMNH) ; 1943 (1♂, 1 sex undetermined CEAH) ; no date (3♀, 2♂ AMNH) ; Curitiba, Vale do Rio Barigui , B. Pohl, 01/I/1943 (1♀ MZSP) . Rio Grande do Sul: Canela, M. Hoffman, 22/ XI/1990 (1♂ DZUP) ; Xanxerê, no collector, XI/1977 (1♂, 11 sex undetermined DZUP) . Santa Catarina: Cauna [Porto União, Santa Cruz do Timbó ], A. Maller, 01/I/1946 (4♀, 7♂ AMNH) ; Corupá, A. Maller, 01/XI/1945 (3♀, 1♂ AMNH) ; 01/XI/1946 (3♀, 2♂ AMNH) ; Florianópolis, B. A. S. Medeiros, 02/XII/2009 (18♀, 14♂ MZSP) ; Imbituba, Areais da Ribanceira , Leonardo K. Sampaio, XI/2010 (2♀, 1♂ MZSP) ; Lages, S. S. Ortolan, XII/1981 (2♀, 1♂ DZUP) ; Laguna, B. A. S. Medeiros, 03/XII/2009 (4♀, 1♂ MACN, 8♀, 45♂ MZSP, 54♀, 4♂ NMNH) ; Nova Teutônia [Seara, Nova Teutônia], Fritz Plaumann, XI/1977 (7♀, 9♂ DZUP) ; Rio Vermelho, Dirings, VIII/1950 (36♀, 37♂ MZSP) ; São Bento do Sul, Dirings, II/1951 (43♀, 33♂ MZSP) . São Paulo: São Paulo, Fabiana D’Agostino, XI/2003 (8♀, 6♂ MZSP) ; São Paulo, Cidade Universitária, B. A. S. Medeiros, 06/X/2009 (2♀, 1♂ MZSP) ; 07/X/2006 (1♀ MZSP) ; 07/X/2009 (1♀ MZSP) ; 18/XI/2009 (1♀ MZSP) .

URUGUAY. Maldonado: Piriápolis , no collector, 15/II/1909 (1♀, 2♂ MLPA) ; Puente l’este [ Punta del Este ], no collector, no date (1♂ MZSP) . Montevideo: Montevideo, J. Tremoleras, no date (1♂ MLPA) ; no collector, no date (1♀ MLPA) ; Peñarol , no collector, 07/II/1909 (1♀ MLPA) ; Prado , no collector, 31/XII/1906 (1♂ MLPA) ; 31/XII/1908 (2♀, 4♂, 1 sex undetermined MLPA) .

NO COUNTRY. No region : No locality, A. Maller, no date (3♀, 2♂ MNRJ) ; A. Mesa, no date (1♂ MLPA) ; C. O., 17/XII/1937 (1♀ MZSP) ; no collector, no date (2♀, 1♂ CEAH) .

ILLEGIBLE. illegible: illegible, Helfer, XI/1960 (1♂ USNM) .

Figures

FIGURE 1. Measurements taken and views used to take them, exemplified with a male of A. aegrotus. Scales: 500 µm. A Head, dorsal view: rostrum width at apex, rostrum width at base, distance between eyes above, head width. B Head, lateral view: rostrum length. C Head, ventral view: distance between eyes below. D Eye, frontal view: eye height and eye length. E Prothorax, dorsal view: prothorax length, prothorax width at base, scutellum length, scutellum width (in case scutellum at a different angle, an additional view with a detail of the scutellum was used). F Prosternum, ventral view: length of prosternum, coxal width. G Elytra, dorsal view: width of one elytrum. H Elytra, lateral view: length of elytra. I Proleg, lateral view: length of profemur, width of profemur, length of protibial, width of protibial. J Antenna, lateral view: length of scape, width of scape, length of each antennomere of the funicle, width of each antennomere of the funicle, length of club, width of club. K Male genitalia, dorsal view: width of aedeagus at base. L Male genitalia, lateral view: Length of aedeagus following the ventral curvature, length of ventral sclerotization beyond the insertion of apodemes, height of aedeagus, length of an apodeme following its curvature.

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 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 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 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.