Research Article |
Corresponding author: Zhiliang Wang ( zlwang@bjfu.edu.cn ) Academic editor: Christopher Majka
© 2016 Xiangyang Lv, Miguel A. Alonso-Zarazaga, Zhishu Xiao, Zhiliang Wang, Runzhi Zhang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lv X, Alonso-Zarazaga MA, Xiao Z, Wang Z, Zhang R (2016) Evemphyron sinense, a new genus and species infesting legume seedpods in China (Coleoptera, Attelabidae, Rhynchitinae). ZooKeys 600: 89-101. https://doi.org/10.3897/zookeys.600.6709
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A new genus Evemphyron Alonso-Zarazaga, Lv & Wang, gen. n., belonging to AttelabidaeRhynchitinae, is described. Its single species, Evemphyron sinense Alonso-Zarazaga, Lv & Wang, sp. n., was reared from larvae found inside seed pods of the legume Callerya dielsiana (Fabaceae, Millettieae) in Sichuan Province (China). The species is figured and placed in the Deporaini because of the presence of minute labial palpi, the strongly crescentic apex of the postmentum, and the apodemes of male IX sternite and female VIII sternite curved sinistro-anterially near their cephalic end. It shows 3-segmented labial palpi and male sex patches on the procoxae, characters that suggest a basal position in the tribe.
Attelabidae , Callerya dielsiana , Deporaini , east Palaearctic, Eusproda , Fabaceae , legume, new genus, new species, Rhynchitinae , systematics
As a part of a long-term project on insect-seed interactions, two of the authors (XYL, ZSX) have been investigating the diversity of insect seed predators of woody trees in a subtropical forest of Dujiangyan City (Sichuan Province, China) since 2002. The Dujiangyan region is in the northern part of the Hengduan Mountains, a biodiversity hotspot and priority area for biodiversity conservation in China. Located in the mountains on the western border of the Sichuan Basin, it is in an ecotone between two biogeographical regions, the Qinghai-Tibetan Plateau and the Chengdu Plain. Climatically, it lies in the middle subtropical zone, characterized by evergreen broad-leaved forests. After checking the weevil specimens obtained, a new species belonging to a new genus was identified. It was thought at first sight to belong to the tribe Rhynchitini (Attelabidae: Rhynchitinae). However, an in depth-study of the available material by another author (MAAZ) revealed that the new species belonged in fact to Deporaini. This weevil species was found to infest seedpods of Callerya dielsiana (Harms) P.K.Lôc ex Z.Wei & Pedley (Fabaceae) (Figs
On 27 October 2013, 262 weevil larvae were collected from seedpods of the plant Callerya dielsiana near Dujiangyan City (Sichuan province). All larvae were placed for adult emergence in a PVC tube (diameter 11 cm, length 40 cm) filled with local soil (30 cm in depth). We checked the emergence of adults once every week from March 2014, and collected adult specimens every day after the emergence of the first adult. In total, nine adult specimens emerged during June, July and September 2014. They were stored in 96% ethanol, and later seven were mounted for the morphological study, leaving two for molecular analysis.
The dry specimens show different degrees of immaturity, mainly affecting their abdomens. Only the male preserved in ethanol was mature enough to allow the extraction of moderately sclerotized genitalia and terminalia. This extraction was done directly in the conserving medium. The abdomen was then soaked overnight in lukewarm 10% sodium hydroxide for digestion of soft tissues. Genitalia and terminalia were photographed in glycerine and later mounted in DMHF (5,5-dimethyl-hydantoin formaldehyde resin) on an acetate card, and pinned together with the tergites and sternites. These have been cross labelled with the specimen in ethanol from which they were extracted.
Descriptions were made using a binocular Nikon SMZ 1500. Photographs (Figures
Original label data have been written below in Chinese script. Added transliterations into pinyin or translations are placed between square brackets. Data from different labels are separated by two slashes (//) and lines within a label by one slash (/).
Nomenclature follows
Evemphyron sinense Alonso-Zarazaga, Lv & Wang, sp. n.
A member of the tribe Deporaini Voss, 1929 as currently understood (cf.
Integument black to brownish, with green to dark bronze metallic shine, some areas on legs, antennae and underside a little lighter.
Vestiture yellow or brown, dense; scales arched to oblique, piliform, those on elytra with apex sometimes flagelliform; brown scales forming a chevron on declivity; scales on dorsum of elytra placed transversally or pointing to outer apical angle of elytra, clearly subparallel to striae only on apical half of 1st interstria, those on anterior half of pronotum, head and metarostrum suberect and pointing forward; scape and funicle with yellow piliform scales, black setae at most as long as scales; tarsomeres densely covered with black piliform scales; very short suberect, arched black setae visible only on apical two thirds of elytral interstriae 9 and 10, rarely visible on other interstriae, and on underside of rostrum.
Mouthparts. (Figs
Rostrum (Figs
Antennae (Fig.
Head moderately elongate, subglobose, very weakly constricted behind eyes in side view, but not in dorsal view. Eyes moderately convex, protruding from head outline, in dorsal view longer than minimum distance between them across forehead, in side view slightly oval.
Pronotum (Figs
Elytra (Figs
Ventral areas. Prosternum short in both sexes, procoxae almost reaching front margin. Hypomera not touching at midline, sternellum large, separating both hypomera and forming part of prothoracic margin (Fig.
Legs. Femora unarmed. Tibiae straight, moderately flattened, moderately widening towards apex, without mucrones or spurs in both sexes; meso- and metatibiae with an outer crenulate (bracteate) keel (
Male genitalia and terminalia. Penis (Figs
Female genitalia and terminalia. Ovipositor with gonoxites very wide in anterior half, and obliquely narrowed to an elongate posterior half (“subdivided” in
The genus name is based on the classical Greek prefix εὖ (well), latinized as ev- (as in Evacanthus) and the present active participle of the verb ἐμφύρω (to confuse), ἐμφῦρον (the confusing one). Gender neuter. Stem is Evemphyront-.
豆毛象属 [dòu máo xiàng shǔ].
Characters as given for the genus. In addition:
Measurements (in mm) (♂, n=5, ♀, n=2): Body length (standard, without head and rostrum): 6.76–7.33. Rostrum: length: 2.83–3.06 (♂), 3.43–3.46 (♀); width (apical): 0.60–0.67. Distance from antennal insertion to base: 1.13–1.33 (♂), 1.47 (♀). Forehead width: 0.53–0.67. Eye length: 0.67–0.73. Scape: 0.28 × 0.16 (♂), 0.28 × 0.12 (♀). Desmomeres: 1: 0.22 × 0.14 (♂), 0.24 × 0.13 (♀); 2: 0.32 × 0.14 (♂), 0.40 × 0.12 (♀); 3: 0.28 × 0.12 (♂), 0.34 × 0.12 (♀); 4: 0.34 × 0.14 (♂), 0.40 × 0.14 (♀); 5: 0.26 × 0.14 (♂), 0.24 × 0.14 (♀); 6: 0.24 × 0.14 (♂), 0.22 × 0.14 (♀); 7: 0.18 × 0.18 (♂), 0.20 × 0.16 (♀). Club: 1: 0.30 × 0.32 (♂), 0.30 × 0.30 (♀); 2: 0.28 × 0.34 (♂,♀); 3: 0.54 × 0.30 (♂), 0.58 × 0.28 (♀). Pronotum: length: 2.33–2.43 (♂), 2.26–2.30 (♀); maximum width: 2.26–2.37. Elytra: length: 4.43–4.90; maximum width: 3.30–3.46.
Rostrum 1.17–1.27 × as long as pronotum in male (Fig.
Antennae inserted at 0.39–0.44 from base of rostrum in male, at 0.42–0.43 in female, articles with integument shining, with moderately long scales and setae, except velvety club, only with a few setae; scape and pedicel oblong, other desmomeres subcylindrical, except 7th, subglobular to suboblong.
Head with forehead forming a very obtuse to almost flat angle with rostrum in side view, forehead densely punctate and pubescent, scales pointing forward, underside of head with strong transverse rugae, these prominent in side view.
Pronotum slightly depressed transversely behind front margin and before hind one, the punctures in the depressions more confused than in the remaining surface.
Elytra 1.33–1.43 × as long as wide, with interstriae densely punctulate, punctures ca. 1/6 the diameter of those forming the striae. Size of strial punctures decreasing towards apex of elytra.
Legs similar in both sexes, devoid of any sexual character.
Wings blackish.
Nine specimens, rather teneral. All printed labels with 中国科学院动物研究所 [Zhōngguó kēxuéyuàn dòngwù yánjiū suǒ, Chinese Academy of Sciences, Institute of Zoology]. All specimens are deposited in this institution, except one male and one female paratype, which are deposited in the Alonso-Zarazaga collection (Museo Nacional de Ciencias Naturales, Madrid, Spain).
Holotype: 1 male, labelled: printed: 2014-VII-31 / 四川 都江堰 浦阳镇 花 / 溪村 [Sìchuān, Dūjiāngyàn, Pŭyángzhèn, Huāxīcūn], 肖治术 [Xiāo Zhìshù] leg. // printed: N31°03'45.07" / E103°43'0.52" / Alt. 709 m // printed: 寄主 香花鸡血藤 [jìzhǔ [host] xiānghuā jī xuè téng] / Callerya dielsiana (Harms) // handwritten: A037 / 1(in red) ♂ / H.7.31.
Paratypes: 1 male, same data as holotype, except dated 2014-VII-12 and a handwritten label: A037, 14-7-12 / 山胡豆[Shānhúdòu], 左后[zuǒhòu] / 足断[zúduàn] ♂; 1 male, same data as holotype, except dated 2014-VII-7 and a handwritten label: A037 / 1(in red) ♂ H.7.7 (specimen had been dissected, the abdomen had been discarded, some pieces in a glycerine vial); 1 male, same data as holotype, except dated 2014-VI-24, and a handwritten label: A037 / H / ♂ 6.24; 1 male, same data as holotype, except dated 2014-VII-7, and a handwritten label: A037 / 2 (in red) / ♂ H.7.7.; 1 female, same data as holotype, except dated 2014-VI-20, and a handwritten label: A037 / ♀ H.6.20; 1 female, same data as holotype, with a handwritten label: A037 / 2 (in red) / ♀ H.7.31 (specimen dissected, abdomen discarded, some pieces in a glycerine vial); 2 males, same data as holotype, but dated 2014-VI-12 and 2014-IX-17 respectively, conserved in pure ethanol vials for DNA extraction. (one of them, being less teneral, has been dissected for the study of the male genitalia and terminalia).
The species is named after the country where it has been found, China (in Latin: sinensis, -e: Chinese). It is an adjective, in neuter form to agree with the gender of the genus.
中华豆毛象 [zhōnghuá dòu máo xiàng].
Callerya dielsiana (Harms) P.K.Lôc ex Z.Wei & Pedley (Fabaceae, Millettieae). Larvae develop in seeds inside the pods. This weevil species was not found on other plant species despite long-term collecting of several Fabaceae species and other possible host plants carried out at the same study site.
This species is known only from the type locality in Sichuan Province (China).
This genus is superficially similar to Cyllorhynchites Voss, 1930, mainly in the head not being constricted behind the eyes and the presence of dense yellowish piliform vestiture, somewhat reminiscent of that of C. (C.) ursulus rostralis (Voss, 1930), a common weevil in China. This led to the initial placement in the tribe “Rhynchitini”, sensu
The relationships of Evemphyron within Deporaini are also contentious. In fact, no genus known to belong to this tribe seems to be closely related. The keys provided by
The new genus should belong, according to the keys proposed by
If the character of the covered propygidium is not taken into consideration, the only possible placement is within the subtribe Deporaina. However, this is also a very disparate group regarding its contents. The combination of 3-segmented labial palpi, propygidium practically covered by the elytra and lacking definite wing folding patches, tibiae neither spurred nor mucronate in both sexes, long rostrum without long, dense patches of long hairs in female and male procoxae with sex patch does not indicate any included genus as an obvious close relative. This set of character states suggests that the new genus could be very primitive (or even the most primitive) in the subtribe, since the palpi segment number, the long rostrum and the male sex patch on the procoxae are clearly symplesiomorphies shared with genera of Byctiscini, Rhynchitini and Auletini, but they are absent from advanced members of Deporaini.
Eusproda shares with Evemphyron the male sex patch on the procoxae (a symplesiomorphy suggesting the basal position of both genera) and the trophic link to Fabaceae, although Eusproda behaves as a shoot cutter of kudzu (Pueraria montana (Lour.) Merr.) and Japanese clover (Lespedeza cyrtobotrya Miq.) (Fabaceae, Phaseoleae and Desmodieae, respectively), and not as an ovary or young fruit driller, as is the case for E. sinense. They also share the long rostrum, the very weak basal constriction of the head, the antennae inserted just behind middle of the rostrum, the non-contiguous hypomera, the subisodiametric pronotum, the absence of a scutellar striole, the tibiae without mucrones or spurs, overall similarity of the male genitalia and the ovipositor divided into two regions and with styli. Table
Character | Eusproda | Evemphyron |
---|---|---|
Integument | black with blue metallic shine mostly on elytra | black to brownish, with green to dark bronze metallic shine, some areas of appendages and abdomen lighter |
Vestiture | sparse, thin, brown | dense, comprised of yellow and brown piliform scales |
Base of rostrum in female | with dense long hairs | without long hairs |
Labial palpi | 2-segmented | 3-segmented |
Second desmomere | about as long as scape or 1st desmomere | longer than scape or 1st desmomere |
Last antennal club segment | as long as 1st or 2nd, symmetrical | longer than 1st or 2nd, asymmetrical |
Eye length in dorsal view | less than forehead width | more than forehead width |
Pronotum | without median keel | with shortened median keel |
Scutellum | oblong | slightly transverse |
Elytra | elongate, ca. 1.7 × as long as wide | shorter, 1.33-1.43 × as long as wide |
Elytra | uniformly convex | dorsally flat to concave behind scutellum |
Elytral striae 9th and 10th | confluent near apex of elytra | confluent at metacoxal level |
Propygidium | with wing folding patches | without wing folding patches |
Metatarsomere 1 | slightly longer than 2+3 | clearly shorter than 2+3 |
Tegminal arms | broad, angulate | thin, curved |
Tegminal manubrium | strongly asymmetrical, T-shaped at apex | slightly asymmetrical, uniformly broadened |
Size | smaller (3.5-4.5 mm, without rostrum) | larger (6.76-7.33, without head and rostrum) |
Biology | shoot-cutter | ovary- and young-fruit- driller |
Evemphyron could be close to any of the genera related to Deporaus Samouelle, 1819. However, the same combination of characters precludes the finding of another genus sharing putative synapomorphic features, as no known genus matches the combination found in Evemphyron. Caenorhinus C.G. Thomson, 1859 shares with Evemphyron the confluence of 9th and 10th elytral stria at metacoxal level and the absence of wing folding patches on the propygidium, but differs clearly by the presence of spurs in at least one pair of tibiae in both sexes and of mucrones at least in one pair of tibiae in males, the shorter, apically widened rostrum and the presence of defined endophallic sclerites.
In summary, this new genus is placed in DeporainiDeporaina on the basis of the minute labial palpi, the strongly crescentic apex of postmentum, the absence of scutellar striole, and the apodemes of male sternite IX and female sternite VIII curved sinistro-anteriad near their cephalic end.
The definitive placement of this new genus will have to wait until a molecular phylogeny of the tribe (and the subfamily Rhynchitinae as a whole) is performed.
We thank Xunlong Wang and Chengqiang Wang for their help with field work. The experiments comply with the current laws of China. This research was supported by National Natural Science Foundation of China (Nos. 31330013, 31270470, 31071929, 31201735, 31172130, 31210103909, J1210002). We are also grateful to Dr Ning Liu and Dr Kuiyan Zhang (Institute of Zoology, Chinese Academy of Science) for their continuous support during the completion of this research. Drs Robert S. Anderson (Canadian Museum of Nature) and Alexander Riedel (Naturkundemuseum Karlsruhe) are warmly acknowledged for their reviews, which improved our work very much. Dr. Chistopher H.C. Lyal (Natural History Museum, London) is warmly thanked for his language correction.