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Los primeros registros de gorgojos (Coleoptera: Curculionidae) del cuaternario colombiano copal.

First records of weevils (Coleoptera: Curculionidae) in Quaternary Colombian copal

Introduction

Copals are Quaternary subfossil resins that can contain both extinct and extant species of animals and plants (Poinar et al. 2001, 2013). Representatives of the superfamily Curculionoidea in copal are known from African, Madagascar, Celebes, Japanese and Danish samples (Legalov 2015) although bark-beetles are the most richly represented group of copal curculionoids.

The importance of copal inclusions is that they can illustrate faunal changes that have occurred relatively recently. Quaternary weevils are mainly represented by recent species (Kiselev and Nazarov 2009). Extinct Pleistocene insects in copal are rare (Hinojosa-Diaz and Engel 2007; Azar et al. 2009; Stroinski and Szwedo 2011; etc.). Of the several hundred species of Late Pleistocene weevils, primarily from river and lake deposits, 24 species (with 16 extant) occurred in copal (Legalov 2015).

In this paper, we describe three new weevil species of the subfamilies Dryophthorinae, Molytinae, and Entiminae from copal of Santander province, Colombia. These are the first records of the family Curculionidae in Colombian copal.

Material and methods

Colombian copal is the subfossil resins of leguminous trees of the genus Hymenaea sp. (Fabaceae) (Langenheim 1995, 2003). The age of Colombian copal still remains an open question, however it is usually dated within the age range of 60 years (postbomb) to 2.5 Ma years [Holocene-Pleistocene-Pliocene] (Anderson 1996; Lambert et al. 1995; Penney et al. 2013; Poinar 1992; Ragazzi et al. 2003; Stinchcomb 1998). Accurate determinations made by means of 14C revealed that some pieces of Colombian copal from the Santander province are approximately 200 years old (Clifford et al. 1997), and this age corresponds to nearly recent resin.

Type strata and locality. Colombian Quaternary copal; Santander province, Andes region, Columbia, South America.

The type specimens are deposited in the Poinar amber collection maintained at Oregon State University (PACO, Corvallis, OR, USA) and the private collection of Andris Bukejs (ABDL, Daugavpils, Latvia).

Observations and photographs were made with a Nikon 140 SMZ-10 stereoscopic microscope, Nikon SMZ 745T stereomicroscope and Zeiss Stemi 2000-C binocular microscope. Helicon Focus Pro X64 and CombineZP were used to stack photos for better clarity and depth of field.

Taxonomy

Curculionidae Latreille, 1802 Dryophthorinae Schoenherr, 1825 Dryophthorini Schoenherr, 1825 Stenommatus Wollaston, 1873 Stenommatus copalicus Poinar, Bukejs et Legalov sp. nov.

(Fig. 1)

Diagnosis. This new species differs from the previous fossil S. pulvereus Davis et Engel, 2006 from Dominican amber in possessing shorter elytra, weaker pronotal punctation, a narrowing of the elytra in the apical and a brown body. It differs from S. leptorhinus Poinar et Legalov, 2015 by its large body and longer pronotum with intervals as wide as the striae. S. copalicus sp. nov. is similar to the recent S. sulcifrons Champion, 1909 from Central America, but differs in its wider body, larger pronotal punctation, and greater convex elytral humeri.

Description. Body elongate, weakly convex dorsally; rufous, head and basal % of rostrum dark brown; glabrous. Head relatively small, 0.6 times as long as rostral length; rostrum quite long, 2.8 times as long as wide in middle, cylindrical, curved, slightly dilated towards apex in apical 1/3; with small and dense punctuation; about 0.8 times as long as pronotum; eyes large, narrowly-oval, almost flat, closely approximate to each other on underside of head; forehead flat, vertex convex; temples large, distinctly longer than transverse diameter of eye; vertex with slight punctuation. Antennae inserted laterally near middle of rostrum, relatively short, reaching anterior margin of pronotum; scape large, elongate, weakly clavate, about 5.5 times longer than wide, 0.6 times as long as rostrum; flagellum short, compact, 4-articled (appears 3-articled), flagellomeres indistinctly visible, subequal in length, trapezoidal, widened distally, last flagellomeres wider than previous; antennal club 1-articled, ovoid with widely rounded apex, dilated apically, nearly as long as flagellum, apically subtruncate. Pronotum elongate, distinctly narrower than elytral base, widest in middle; with deep constriction in anterior anterior and posterior margins straight, lateral margins weakly rounded, anterior margin narrower than posterior margin; pronotal punctures round, dense and large (slightly smaller than elytral punctures); distance between adjacent punctures distinctly smaller than diameter of puncture; intervals convex; scutellum minute, subtriangular. Elytra ovoidelongate, weakly convex, widest in basal 1/3; humeri convex; base with convex borders; subparallel in basal V2, in apical 1/4 abruptly narrowed towards apex and jointly rounded apically, about 1.8 times as long as pronotum; elytral punctures round, large and dense, arranged in regular striae; distance between strial punctures convex, very narrow, about 4.05.0 times smaller than diameter of puncture; striae distinct throughout entire length of elytra; intervals very narrow (as distance between strial punctures) and convex; pygidium not exposed. Legs relatively short; femora weakly clavate, slightly flattened, without denticles; tibiae flattened, straight with parallel sides; uncus large with sickle-shaped inward curve; mucro tooth-shaped, small; apical setal combs absent; tarsi 5-jointed, slender, short, about 1/2 as long as tibiae; 1st-4th tarsomeres trapezoidal, 1st-2nd tarsomeres small, nearly equal, about as wide as uncus at base; onychium elongate, subcylindrical, 0.9 times as long as tarsomeres 1st-4th combined; claws small, free and simple.

Measurements. Body length (without rostrum) 2.2 mm, rostrum length (in lateral view) 0.45 mm.

Etymology. The specific epithet of this new species is formed from the word "copal".

Material examined. Holotype: "Nr. AB 035" [white printed label], "Holotype / Stenommatus copalicus sp. nov." [red printed label] in abdl; sex unknown. A complete beetle embedded in an elongate copal piece (length 42 mm, width 21 mm); ventral side of specimen almost invisible because of its location in copal piece. Animal syninclusions: one winged Isoptera; 6 undetermined small insects; few body parts of insects (antenna, legs, tarsus). Few very small gas bubbles and little cracks diffusely spread throughout the copal piece.

Remarks. This new species belongs to the subfamily Dryophthorinae based on its geniculate antennae with a compact antennal club, 1st article of club elongated with other articles fused, pronotum without lateral carina, and flagellum with 4 flagellomeres. The flagellum with 4 flagellomeres, 5-jointed tarsi, antennae inserted near the middle of the rostrum, and pygidium exposed indicate placement in the tribe Dryophthorini. The rostrum almost equal in length to the pronotum, the eyes closely approximate to each other on the underside of the head, and an apically subtruncate antennal club are characteristics of the genus Stenommatus.

Key to recent and fossil species of Stenommatus in New World
1. Pronotum finely punctate; elytral intervals 2.0-2.5 times
As long as striae width. Bolivia S. inflexus Hustache, 1938

1 'Pronotum coarsely punctate; elytral intervals as wide as or
Narrower than striae width                                       2

2. Rostrum elongate, 4.5 times as long as wide in middle.
Dominican amber S. tanyrhinus Poinar et Legalov, 2015

2' Rostrum short, 1.7-3.0 times as long as wide in middle        3

3. Rostrum thick, 1.7-1.8 times as long as wide in middle        4

3' Rostrum thin, 2.8-3.0 times as long as wide in middle         6

4. Body wide. Mexico S. fryi
Wollason, 1873

4' Body narrow                                                   5

5. Antennal scrobe very shallow; body black. Dominican
amber S. pulvereus Davis et Engel, 2006

5' Antennal scrobe distinct; body black, but rostrum, scape,
flagellum, tarsi brownish. Central America S. sulcifrons
Champion, 1909

6. Body large (2.2 mm); pronotum longish; elytra 1.9 times
As long as pronotum; striae narrow, with intervals as wide as
striae. Colombian copal S. copalicus sp. nov.

6' Body small (1.4 mm); pronotum shortish; elytra 2.4 times
as long as pronotum; striae wide, with intervals 0.4-0.6
times as wide as striae. Dominican amber S. leptorhinus
Poinar et Legalov, 2015.


Molytinae Schoenherr, 1823 Conotrachelini Jekel, 1865 Conotrachelus Dejean, 1835 Conotrachelus dysaethrius Poinar, Bukejs et Legalov, sp. nov. (Fig. 2)

Diagnosis. This new species differs from C. pertusicossis Fiedler, 1944 from Colombia by its rufous body, rostrum slightly curved, scutellum rhombic and apex of elytra not rounded. From the Colombian C. subquadraticollis Fiedler, 1944, it differs by the longer rostrum, narrower pronotum, rhombic scutellum, smooth humeri, and the apex of the elytra bearing large blunt teeth.

Description. Body slightly elongate, convex dorsally; rufous; glabrous; with erect and semierect quite elongate scales. Rostrum long, slightly curved; small and densely punctate; 12.1 times longer than width at apex, 9.4 times longer than width in middle, 5.7 times longer than width at base; 1.1 times as long as pronotum; eyes large, oval, almost flat; forehead almost flat; vertex weakly convex, densely punctate; temples short. Antennae inserted laterally before middle of rostrum, quite long; scape large, elongate; antennal club compact. Pronotum bell-shaped, without middle carina and middle stria, without erect groups of scales; 1.5 times longer than width at apex, 0.7 times longer than width in middle and at base, widest near middle, narrowing to apex; lateral margins almost straight; anterior margin 0.5 times as long as posterior margin; pronotal punctures round, dense and large; distance between adjacent punctures distinctly smaller than diameter of puncture; intervals convex; scutellum rhombic, 1.5 times longer than width in middle. Elytra almost trapezoidal, convex, widest in basal 1/3, 1.6 times longer than wide at base and in middle, 2.4 times longer than wide at apical fourth, 2.5 times as long as pronotum; in basal quarter subparallel, further narrowing towards apex; elytral apex truncated with toothed portions; humeri smooth; elytral striae regular and distinct; intervals carinate, 4.0 times as wide as striae; carinae of 2nd and 5th intervals broken. Prosternum with distinct postocular lobes; precoxal portion of prosternum elongate; postcoxal portion of prosternum short; procoxal cavities separated; prosternum with ventral channel; mesocoxal cavities widely separated; mesepimeron not ascended; metasternum quite short; metepisternum quite broad, 6.1 times longer than width in middle. Abdomen flattened; 1st ventrite elongate; 2nd ventrite 0.7 times as long as 1st ventrite; 2nd--4th ventrites almost equal in length; 5th ventrite 1.3 times as long as 4th ventrite; pygidium not exposed. Legs quite long; femora distinctly clavate, slightly flattened, large tooth; tibiae flattened, curved, with apical setal combs and ancus, without mucro; uncus large and curved; mesotibiae 4.5 times longer than width in middle; metatibiae 4.7 times longer than width in middle; tarsi long; 1st-3rd tarsomeres conical; 5th tarsomere elongate; tarsomeres with pulvilli on underside; claws free, large, clearly diverging, with teeth.

Measurements. Body length (without rostrum)--8.5 mm, rostrum length--2.6 mm.

Etymology. From the Greek "dysaithrios" = murky Material examined. Holotype: C-7-345A in PACO.

Remarks. This new species belongs to the subfamily Molytinae based on the long tibiae with large unci and the mesepimeron not being ascended. The prosternum with a ventral channel suggests placement in the tribe Conotrachelini. The pronotum with distinct postocular lobes and elytra with carinate intervals indicates placement in the genus Conotrachelus and to sub group 2 of group 2 (sensu Fiedler 1944) based on the broken carinae of the 2nd and 5th intervals.
Key to recent and fossil species of Conotrachelus with carinae
of 2nd and 5th elytral intervals broken and body
size greater than 7.0 mm

1. Rostrum shorter than head and pronotum together. Columbia
C. subquadraticollis Fiedler, 1944

1' Rostrum as long as head and pronotum together                   2

2. Body rufous; rostrum slightly curved; scutellum rhombic;
apex of elytra containing large blunt teeth. Columbian copal
C. dysaethrius sp. nov.

2' Body black; rostrum distinctly curved; scutellum trapezoidal;
apex of elytra rounded. Columbia C. pertusicossis
Fiedler, 1944.


Entiminae Schoenherr, 1823 Entimini Schoenherr, 1823 Cydianerus Schoenherr, 1840 Cydianerus eukrinus Poinar, Bukejs et Legalov, sp. nov.

(Fig. 3)

Diagnosis. This new species differs from C. latruncularius (Perty, 1832) from South America by the carinate elytral intervals, elytra parallel-sided, a narrower, granulated pronotum not constricted to the base and the antennae not reaching the humeri. From C. araneiformis (Dalman, 1833) from Brazil it differs by the granular pronotum, narrow carina of elytral intervals, elytra parallel-sided, longer rostrum and antennae not reaching the humeri. From C. flexuosus (Pascoe, 1881) from Brazil it differs by the granular pronotum, unicolorous scales on the elytra and carinate elytral intervals.

Description. Body wide-elongate, convex dorsally; rufous; shiny. Head large, 0.4 times as long as rostral length; rostrum short, curved, dilated towards the apex in apical 1/3; large and densely punctate; 0.9 times as long as pronotum; with 3 carinae; nasal plate prominent; mentum with setae; scrobes oblique; pterygium partially visible from above; eyes large, oval, weakly convex, 0.7 times longer than wide; forehead flat, 0.8 times as wide as rostrum base width; temples short, 0.4 times as long as eye length; vertex weakly convex, densely punctate; mandibles with prominent scars of attachment of deciduous process. Antennae inserted laterally near apical third of rostrum, elongate, almost reaching anterior margin of elytra base; scape large, elongate, 4.5 times longer than wide, 0.7 times as long as rostrum, reaching eye; flagellum long, 7-articled; flagellomeres trapezoidal, widened distally; 1st flagellomere 2.0 times longer than wide, 0.3 times as long and 0.6 times as wide as scape; 2nd flagellomere 1.8 times longer than wide, 0.7 times as long and 0.8 times as wide as 1st flagellomere; 3rd flagellomere 1.6 times longer than wide, 0.6 times as long and 0.8 times as wide as 2nd flagellomere; 4th flagellomere 1.4 times longer than wide, equal in length and 1.1 times as wide as 3rd flagellomere; 5th flagellomere 1.1 times longer than wide, 1.1 times as long as and 1.4 times as wide as 6th flagellomere; 6th flagellomere 1.6 times longer than wide, 1.4 times as long as and 1.4 times as wide as 5th flagellomere; 7th flagellomere 1.4 times longer than wide, equal in length and 1.1 times as wide as 6th flagellomere; antennal club 3-articled, 0.5 times as long as flagellum; 1st club article 1.5 times longer than wide, 1.6 times as long and 1.5 times as wide as 7th flagellomere; 2nd club article 0.6 times longer than wide, 0.5 times as long as and 1.2 times as wide as 1st club article; 3rd club article 1.3 times longer than wide, 1.7 times as long as and 0.9 times as wide as 2nd club article. Pronotum bell-shaped, 1.1 times longer than wide at apex, 0.8 times longer than wide in middle, 0.9 times longer than wide at base, distinctly narrower than elytral base; greatest width before middle; disk weakly convex, with large granulation; distance between granules distinctly smaller than diameter of puncture; scutellum trapezoidal, convex. Elytra convex, 2.2 times longer than wide at base, 1.5 times longer than wide at humeri, 1.4 times longer than wide in middle, 1.7 times longer than wide at apical fourth; 2.8 times as long as pronotum; elytral base 1.1 times as wide as pronotum base; greatest width before middle; humeri weakly convex; elytral punctures round, large and dense, arranged in regular striae; distance between strial punctures convex, very narrow, about 4.0-5.0 times smaller than diameter of puncture; striae distinct throughout entire length of elytra; intervals very narrow (as distance between strial punctures) and convex; pygidium not exposed. Thorax punctate; precoxal portion of prosternum equal in length to postcoxal portion, procoxal cavities contiguous, 1.8 times as long as precoxal portion; postocular lobes of prosternum with long vibrissae mesocoxal cavities widely separated, metasternum tumid; metepisternum quite narrow. Abdomen convex ventrally; 1st and 2nd ventrites elongate; 1st ventrite shorter than 2nd and 3rd ventrites combined; 2nd ventrite 0.7 times as long as 1st ventrite; 3rd ventrite 0.8 times as long as 2nd ventrite; 4th ventrite 0.6 times as long as 3rd ventrite. Legs long; femora weakly clavate, without denticles; profemora 2.4 times longer than wide; mesofemora 3.4 times longer than wide; metafemora 3.6 times longer than wide; tibiae flattened, almost straight, without uncus and mucro, with apical setal combs; protibiae 7.3 times longer than wide in middle; mesotibiae 8.6 times longer than wide in middle; metatibiae 8.6 times longer than wide in middle; tarsi long and quite wide; 1st and 2nd tarsomeres trapezoidal; 3rd tarsomere bilobed; onychium elongate, subcylindrical; claws long, simple, connate at base; metatarsus: 1st tarsomere 2.2 times longer than wide at apex; 2nd tarsomere 1.3 times longer than wide at apex, 0.8 times as long and 1.3 times as wide as 1st tarsomere; 3rd tarsomere bilobed, 0.7 times longer than wide at apex, equal in length and 1.8 times as wide as 2nd tarsomere; 5th tarsomere 3.0 times longer than wide at apex, 1.2 times as long and 0.3 times as wide as 3rd tarsomere.

Measurements. Body length (without rostrum)--6.7 mm, rostrum length--3.0 mm.

Etymology. From the Greek "eukrines" = clear

Material examined. Holotype: C-7-345B in PACO. Remarks. This new species belongs to the subfamily Entiminae based on the mandibles with prominent scars of attachment of deciduous processes, rostrum short and tibiae without uncus. The swollen metasternum and 2nd ventrite shorter than the 3rd and 4th ventrites combined suggest placement in the tribe Entimini. The prominent nasal plate, protibiae without mucro, 1st ventrite shorter than 2nd and 3rd ventrites combined; tarsal claws connate at base, scrobes oblique, mentum with setae and postocular lobes with long vibrissae indicate placement in the genus Cydianerus.
Key to recent and fossil species of Cydianerus in New
World

1. Pronotum granulated                                            2

1' Pronotum punctate                                              4

2. Elytral intervals carinate. Elytra parallel-sided, narrower.
Pronotum granulated, not constricted at base. Antennae
not reaching humeri. Colombian copal C. eukrinus
sp. nov.

2' Elytral intervals convex. Elytra narrowed from humeri to
apex, wider. Pronotum wrinkly-granulated, constricted at
base. Antennae reaching humeri                                    3

3. Body with green scales. Brazil.... C. virescens Lucas, 1857
3' Body with light scales. South America C. latruncu
larius (Perty, 1832)

4. Elytral intervals carinate. Brazil C. araneiformis
(Dalman, 1833) and C. bohemani Boheman, 1840

4' Elytral intervals convex                                       5

5. Elytra yellow with dark spots                                  6

5' Elytra not yellow, dark-color, or with transverse fasciae
7

6. Elytra parallel-sided. Brazil C. ornatus (Pascoe, 1881)

6' Elytra narrowed from humeri to apex. Venezuela
C. wagneri Bovie, 1907

7. Elytra with transverse fasciae. South America
C. flexuosus (Pascoe, 1881) and C. bifasciatus
Hustache, 1926

7' Elytra with unicolorous scales                                 8

8. Elytra wide, with dark blue scales. Panama, Colombia
C. pascoei (Bovie, 1908)

8' Elytra narrower, with silvery scales. Central America
C. argenteus Boheman, 1840


Discussion

The genus Stenommatus Wollaston comprises more than 10 species distributed in Central and South America, Madagascar, Mascarene Is., Society Is., India, Japan, New Guinea and Australia (Hustache 1924; Osella 1979; Alonso-Zarazaga and Lyal 1999). Three species were described from Miocene Dominican amber (Davis and Engel 2006; Poinar and Legalov 2015). The beetles probably occur under the bark of dead or dying trees (Morimoto 1978). The biology of Dryophthorini is poorly investigated.

Conotrachelus is the largest genus of the subfamily Molytinae distributed in the New World and numbers more than

1090 species (Fiedler 1940, 1944; Wibmer and O'Brien 1986; O'Brien and Wibmer 1982). New species have been described fairly recently (O'Brien and Couturier 1995; Salas-Araiza and Romero-Napoles 2012). The beetles are associated with various plants and the larvae develop in fruits or galls (Schoof 1942). Only C. florissantensis Wickham, 1912 from the Late Eocene of the Florissant beds in North America is known in fossil form (Wickham 1912).

The tribe Entimini is represented by 7 genera with 44 species in Central and South America, namely Rhigus Schoenherr, 1823 (13 species), Cydianerus (10 species), Polyteles Germar, 1829 (7 species), Entimus Germar, 1817 (7 species), Phaedropus Schoenherr, 1823 (5 species), Nesocomptus Vanin et Gaiger, 2005 (1 species) and Trachyus Kuschel, 1955 (1 species) (Gaiger 2001; Morrone 2002; Vanin 1983; Vanin and Gaiger 2005; Vaurie 1952). One species from the genus Entimus was described from the Middle Eocene Green River (USA) deposits (Scudder, 1876).

Received: 27-Jan-2016 * Accepted: 26-Jul-2016

Suggested citation:

POINAR, G. JR.; BUKEJS, A.; LEGALOV, A. A. 2017. First records of weevils (Coleoptera: Curculionidae) in Quaternary Colombian copal. Revista Colombiana de Entomologia 43 (1): 85-90. Enero-Junio 2017. ISSN 0120-0488.

Acknowledgements

We thank Alex E. Brown (California: Berkeley) for providing the Conotrachelus fossil and are grateful to O. Jaeger (Germany: Dresden), K.-D. Klass (Germany: Dresden), P. Limbourg (Belgium: Brussels), H. Perrin (France: Paris), A.G. Kirejtshuk (Russia: St.-Petersburg and France: Paris), B.A. Korotyaev (Russia: St.-Petersburg) for the opportunity to study comparative material deposited in the Institut Royal des Sciences Naturelles de Belgique, Museum National d'Histoire Naturelle (France: Paris), Museum fur Tierkunde, Senckenberg Naturhistorischen Sammlungen Dresden and the Zoological Institute of the Russian Academy of Sciences.

The study was partially supported by the Federal Fundamental Scientific Research Programme for 2013-2020, project no. VI.51.1.7 and the Russian Foundation for Basic Research, project no. 15-04-02971a.

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George Poinar, Jr. (1), Andris Bukejs (2), Andrei A. Legalov (3)

(1) Ph. D. Department of Integrative Biology, Oregon State University, Corvallis, OR 97331 USA, poinarg@science.oregonstate.edu, corresponding author.

(2) Ph. D. Institute of Life Sciences and Technologies, Daugavpils University, Vienibas Str. 13, Daugavpils, Latvia, carabidae@inbox.lv. 3 Ph. D., Doctor of Science. Institute of Systematics and Ecology of Animals, ul. Frunze 11, Novosibirsk, 630091 Russia, fossilweevils@gmail.com.

Leyenda: Figure 1. Stenommatus copalicus sp. nov., holotype, habitus, dorsolateral view.

Leyenda: Figure 2. Conotrachelus dysaethrius sp. nov., holotype, habitus: A. Dorsal view. B. Ventral view. C. Lateral view.

Leyenda: Figure 3. Cydianerus eukrinus sp. nov., holotype, habitus: A. Dorsal view. B. Ventral view. C. Lateral view.
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Author:Poinar, George Jr.; Bukejs, Andris; Legalov, Andrei A.
Publication:Revista Colombiana de Entomologia
Date:Jan 1, 2017
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