Phorocardius manuleatus ( Candeze , 1888)

Ruan, Yongying, Douglas, Hume B., Qiu, Lu, Chen, Xiaoqin & Jiang, Shihong, 2020, Revision of Chinese Phorocardius species (Coleoptera, Elateridae, Cardiophorinae), ZooKeys 993, pp. 47-120 : 47

5. Phorocardius manuleatus ( Candeze, 1888) Figs 10 , 11 , 23E , 24E , 25E , 26E

Cardiophorus manuleatus Candèze, 1888: 681. Type locality: “Thagatà, Tenasserim", interpreted as Myanmar, Kayin State, mountains east of Kyaikdon using Hallermann et al. (2002). Lectotype designated here.

Phorocardius melanopterus manuleatus : Fleutiaux 1931: 311.

Phorocardius manuleatus : Fleutiaux 1947: 366.

Differential diagnosis.

Body length greater than 7.0 mm; integument shiny, black with yellow in most, entirely black to black-brown in some. Prothorax: procoxal cavities open; prosternal process strongly narrowed posterad to ventral apex in ventral view, with apex acute. Pterothorax: scutellar shield with posterior apex pointed. Tarsal claw with ventral apex not smaller than dorsal apex. Male genitalia: paramere with preapical lateral expansion present, without apical mesal callus. Female: apex of last abdominal ventrite (ventrite V) simple, not emarginate at apex.

Phorocardius manuleatus ( Candèze, 1888) is unique among Chinese Phorocardius species for its variable color pattern. Some individuals are entirely black to black-brown throughout body, resembling P. yanagiharae (Miwa, 1927) and P. yunnanensis sp. nov.

This species can be differentiated from P. yanagiharae by the following combination of characters. In P. manuleatus : in ventral view, parameres of aedeagus with sides gently narrowed from mid-length to apex (not abruptly narrowed from apical third to near apex), with width 1.5-2 × that of median lobe (measured at apical fourth); and in dorsal view, pronotum with lateral sides of posterior angles almost straight, slightly convex (bulged) at posterior half in a few cases (e.g., in Fig. 11A ); while in P. yanagiharae , in ventral view, paramere of aedeagus with sides abruptly narrowed from apical third to near apex, with width 2-3 × that of median lobe (measured at apical fourth); and in dorsal view, pronotum with lateral sides of posterior angles strongly bulged and convex (Fig. 18C, D ).

This species can be differentiated from P. yunnanensis Ruan & Douglas, sp. nov. by the following combination of characters. In P. manuleatus , in ventral view, paramere of aedeagus narrow and slender near apex, with width to that of median lobe ratio 0.5-0.7 (measured at the area posterior of preapical lateral expansion); legs darker in apical half, not unicolor yellow-brown to brown; while in P. yunnanensis Ruan & Douglas, sp. nov., in ventral view, paramere of aedeagus wide and strong near apex, with width to that of median lobe ratio 1.0-1.2 (measured at the area posterior of preapical lateral expansion); and legs unicolor, entirely yellow-brown.

Distribution.

China (Yunnan, new record), Myanmar ( Candèze 1888), Laos ( Fleutiaux 1931, 1947), Vietnam ( Fleutiaux 1918, 1947).

Description.

(based on lectotype and 24 non-type specimens) Integument shiny, black with yellow in most, or entirely black to black-brown. Pronotum entirely black or orange with variable median black stripe (Fig. 11E ). Ventral side of prothorax yellow, orange, black or orange with black prosternum. Elytra black, yellow, or black with orange spot at elytral bases. Mesosternum brown to black. Metasternum yellow to black. Abdominal ventrites yellow, black, or bicolored (I-IV orange, V yellow). Head red-brown to black. Antennae brown. Legs variably orange to yellow-brown from coxa to mid tibia, yellow-brown to brown from mid tibia to last tarsomere. Body with yellow pubescence.

Measurements.

(based on lectotype and examined specimens) Male body length 7.2-9.6 mm, width 2.2-2.6 mm. Female body length 8.5-9.7 mm, width 2.5-2.9 mm. Body length to width ratio 3.0-3.1. Pronotal width to length ratio 1.1-1.2. Pronotum narrower than elytra, pronotal width to body width ratio 0.87-0.90. Elytral length to pronotal length ratio 2.4-2.6; elytron length to width ratio 4.1-4.2.

Head. Frons and vertex with interspaces between punctures 2.5-6 × average diameter of puncture (Fig. 11D ). Frontal carina in frontal view convex, not straight. Antenna with apex extending to posterior angle of pronotum. Distance between eyes to width of eye ratio 2.7-3.1. Antenna length to body length ratio, in male 0.41-0.42, in female 0.39-0.40. Proportions of antennomere lengths (male): 100 (scape); 51-55; 72-80; 75-82; 82-88; 82-84; 72-78; 72-78; 80-88; 80-89; 114-120.

Prothorax. Pronotum in dorsal view: sides convex near mid-length, nearly straight at ends, widest near mid-length; posterior angles with lateral sides almost straight, slightly convex (bulged) at basal half in a few cases (e.g., Fig. 11A ); surface with interspaces between punctures 4-8 × average puncture diameter (Fig. 11C ). In ventral view, ventral surface of prosternal process with sides carinate and strongly narrowed from anterior to posterior end, with apex acute. In lateral view, prosternal process with ventral surface curved slightly dorsad, posterior end strongly concave (Fig. 24E , upper arrow). Procoxal cavities open.

Pterothorax (Figs 24E , 25E ). Mesepisternum in ventral view with antero-mesal corner angulate mesad of a notch (Fig. 25E , upper (green) arrow). Projections on posterior edge of mesosternum: in ventral view present (Fig. 25E , lower (red) arrow); in lateral view present, acute, strongly produced anteriorly (Fig. 24E , lower (red) arrow). Scutellar shield: width to length ratio 1.0, anterolateral edges slightly sinuate, posterior apex pointed. Elytra: upper edge of epipleura with minute serrations.

Legs. Length ratio of metatarsomeres I-V: 100; 82-92; 67-75; 60-70; 155-180. Claw with ventral apex almost as large as dorsal apex.

Abdomen. Serrations on lateral edges of visible abdominal ventrites I-V absent.

Male genitalia. Robust in ventral view, slender in lateral view. Median lobe in ventral view gradually narrowing from base to near apex, then dilated to rounded apex. Median lobe in lateral view curved ventrad at base, straight from basal third to apex; apex broadly rounded. Paramere in ventral view: robust, width 3-4 × median lobe width (measured at mid-length of paramere and median lobe respectively), widest near mid-length; apical fourth gradually narrowing towards apex, with mesal side bent and turned ventrad in varying degree, result in slightly different shapes in ventral view; apex of paramere slender and sharp, with preapical lateral expansion acute, hook-like to rounded, facing laterally (Figs 10F , 11F , indicated by blue arrow), without apical mesal callus. Paramere in lateral view: slender, almost straight from base to mid-length, curved ventrad and gradually narrowed from mid-length to apex; apex obliquely truncate; preapical ventral expansion acute but not hook-like (Figs 10E , 11F , indicated by blue arrows).

Female. Body color like male. Apex of abdominal ventrite V convex, somewhat angulate (Fig. 26E ). Proximal sclerites of bursa copulatrix ovoid-triangular shaped (Fig. 10I ), basal edge almost without concavity: each with 9-11 large spines mainly on the convex mesal edge, 15-20 smaller spines on disc.

Type material.

Lectotype. ♂ (RBINS): 1) Coll. R. I. SC. N. B., Inde; 2) Tenasserim, Thagatà, Fea. Apr. 1887; 3) Collection E. Candèze; 4) Manuleatus cdz., Tenasserim; 5) Cardiophorus manuleatus , Cd., dèt. E. Candèze; 6) Probably syntype var. a., Det. W. Suzuki, 1986; 7) Lectotype, Cardiophorus manuleatus Candèze, 1888, Des. Ruan & Douglas, 2020.

Additional material.

1♂ (NHMUK), labels: 1) Cotype; 2) Carin Chebà, 900-1100 m, L. Fea, V XII-88; 3) Andrewes Bequest. B. M. 1922-221. 4) Cardiophorus manuleatus Cand. Co.type.; 5) Not paratype of manuleatus Cand., wrong loc., C.M.F. von Hayek. det., 1957. [Notes: locality of this specimen ( “Chebà”) differs from what Candèze (1888) provided ( “Thagatà, Tenasserim"). Although “Thagatà” and “Chebà” are both in "Carin State" (Now Kayin State), Myanmar and there is a ‘Cotype’ label under the specimen, it is still unknown if this specimen belongs to the syntypes described by the author.]

3♂1♀ (IZCAS), labels: 1) Yunnan, Xi-shuang-ban-na, Meng-a (勐阿), 1050-1080 m, Chinese Academy of Sciences [in Chinese]; 2) 1958.VI.2-10, leg. Shuyong Wang [in Chinese]; 3) Phorocardius flavus Det. Shihong Jiang, 1999; 4) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂ (IZCAS), labels: 1) Yunnan, Xi-shuang-ban-na, Meng-zhe (勐遮), 1200 m, Chinese Academy of Sciences [in Chinese]; 2) 1958.IV.14, leg. Shuyong Wang [in Chinese]; 3) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂ (IZCAS), labels: 1) Da-nuo-you IV B 26.04.2009 leg. L.Z.Meng, gift from Na-ban-he Nature reserve [in Chinese]; 2) Yunnan, Jing-hong, Na-ban-he Nature reserve, Meng-song county, Da-nuo-you (大糯有), 2009.IV.26, 770 m, Chinese Academy of Sciences [in Chinese]; 3) 22.20699°N, 100.63761°E, Malaise trap, leg. Linzeng Meng, Chinese Academy of Sciences [in Chinese]; 4) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♀ (SZPT), labels: 1) Yunnan, Xi-shuang-ban-na, Meng-hun (勐混), Chinese Academy of Sciences [in Chinese]; 2) 1958.V.31, leg. Chun-pei Hong [in Chinese]; 3) Phorocardius flavus Det. Shihong Jiang, 1999; 4) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♀ (IZCAS), labels: 1) Yunnan, Xi-shuang-ban-na, Meng-hun (勐混), Chinese Academy of Sciences [in Chinese]; 2) 1958.VI.12, leg. Yirang Zhang [in Chinese]; 3) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂ (IZCAS), labels: 1) Yunnan, Xi-shuang-ban-na, Da-meng-long (大勐龙), 650 m, Chinese Academy of Sciences [in Chinese]; 2) 1958.IV.18, leg. Fu-ji Pu [in Chinese]; 3) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂ (IZCAS), labels: 1) Yunnan, Xi-shuang-ban-na, Meng-la (勐腊), 620-650 m, Chinese Academy of Sciences [in Chinese]; 2) 1958.V.17, leg. Fa-cai Zhang [in Chinese]; 3) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 4♂2♀ (IZCAS), labels: 1) Yunnan, Meng-la (勐腊), 670 m, Chinese Academy of Sciences [in Chinese]; 2) 1982.IV.20, leg. Subai Liao [in Chinese]; 3) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 3♂1♀ (IZCAS), labels: 1) Yunnan, Gan-lang-ba (橄榄坝), 560 m, Chinese Academy of Sciences [in Chinese]; 2) 1957.IV.19, leg. Guangji Hong [in Chinese]; 3) Phorocardius flavus Det. Shihong Jiang, 1999; 4) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂1♀ (IZCAS), labels: 1) Yunnan, Si-mao (思茅), 1200 m, 1957.V.11, leg. Shuyong Wang [in Chinese]; 2) leg. Guangji Hong [in Chinese]; 3) Phorocardius flavus Det. Shihong Jiang, 1999; 4) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018. • 1♂ (IZCAS), labels: 1) Yunnan, close to Si-mao (思茅), 750 m, 1957.V.11, leg. Д. панфилов [Russian name, written in Chinese]; 2) Phorocardius manuleatus ( Candèze, 1888) Det. Ruan, 2018.

Remarks.

This species is unusual for its extensively varied body color. The following three main patterns were found in examined specimens. Color pattern 1 (Fig. 10A, C ): black on dorsum, with orange or yellow spot on base of each elytron; venter black before mesocoxae, orange from mesocoxae to abdominal ventrite IV, brown-black on abdominal ventrite V; antenna brown; and leg orange, yellow from coxa to mid-length of tibia in some, brown from mid-length of tibia to apex. Color pattern 2 (Fig. 11E ): head black; pronotum with different combinations and proportions of colors, orange at sides in most, black near midline and posterior; elytron black, yellow or mixed with brown and yellow; venter mixed with orange and black; and leg orange or yellow from coxa to mid-length of tibia, brown from mid-length of tibia to apex. Color pattern 3: dorsum and venter entirely black to black-brown, antennae brown, and legs yellow-brown on basal half, brown on apical half.

The aedeagus slightly varied in the apex shapes of the median lobe and parameres. In rare cases, the sides of the pronotum are dramatically wider and more robust (e.g., Fig. 11E 1). A comparison of specimens using mitochondrial DNA would be useful to test species boundaries.

Based on specimens from Yunnan, this species inhabits low to middle elevations (ca. 500-1200 m). Yunnan is mountainous, rainy, subtropical to tropical, with subtropical evergreen broad-leaf forest or tropical rain forest. Some of our specimens are collected from Xi-shuang-ban-na tropical rain forest. Known from Oriental Region only.

This species was treated as a subspecies of Phorocardius melanopterus ( Candèze, 1878) by Fleutiaux (1931: 311). We have studied the photograph of the single type specimen of P. melanopterus [RBINS, label information: Coll. R. I. SC. N. B., CAMBODGE // Collection E. Candèze // n. sp. Melanopterus cdz., Cambodia // Cardiophorus melanopterus , Cd., dèt. E. Candèze // Type]. In that specimen, the head is brown-black, the rest of the body is entirely brown throughout including legs and basal four antennomeres (all other antennomeres are missing on the type specimen), and the pronotum with lateral carina diverging from hind angle carina. Its color is different from all known color patterns of P. manuleatus . Additionally, P. melanopterus probably does not belong to Phorocardius because of the presence of pronotal lateral carina (see checklist above).

Figures

Figure 10. Phorocardius manuleatus Candeze, 1888. A male habitus, dorsal view B male habitus, ventral view, arrow indicating claws C female habitus, dorsal view D aedeagus, ventral view (immersed in glycerin) E aedeagus, lateral view, arrow indicating apex of paramere F aedeagus, dorsal view (immersed in glycerin), arrow indicating apex of paramere G ovipositor, dorsal view H female abdominal sternite VIII, dorsal view (near type locality) I distal (top) and proximal sclerites of bursa copulatrix.

Figure 11. Phorocardius manuleatus Candeze (1888). A pronotum and scutellar shield, dorsal view B head, frontal view C punctation on disc of pronotum D punctation on vertex of head E color variation between four individuals; insets 2, 4, 6, 8 are the ventral views of 1, 3, 5, 7 F aedeagus, in dorsal view and lateral view, arrows indicating apices of parameres.

Figure 18. Phorocardius yanagiharae (Miwa, 1927). A holotype, pronotum, dorsal view B holotype, head, frontal view C right posterior angle of pronotum, dorsal view D left posterior angle of pronotum, dorsal view.

Figure 23. Shape of supraantennal carina of Chinese Phorocardius species. Arrows indicating supraantennal carina forked near eye. A P. alterlineatus Ruan & Douglas, sp. nov. B P. flavistriolatus Ruan & Douglas, sp. nov. C P. florentini. D P. magnus. E P. manuleatus. F P. minutus Ruan & Douglas, sp. nov. G P. rufiposterus Ruan & Douglas, sp. nov. H P. unguicularis. I P. yunnanensis Ruan & Douglas, sp. nov. J P. zhiweii Ruan, Douglas & Qiu, sp. nov. K head, frontal-lateral view, arrows indicating forked supraantennal carina.

Figure 24. Lateral view of prosternal process and mesosternum of Chinese Phorocardius species. Upper green arrows indicating shape of posterior end of prosternal process; lower red arrows indicating projections (absent or present) on posterior edge of mesosternum (i. e., on anteroventral angle of mesosternal fossa according to Douglas 2003). A P. alterlineatus Ruan & Douglas, sp. nov. B P. flavistriolatus Ruan & Douglas, sp. nov. C P. florentini. D P. magnus. E P. manuleatus. F P. minutus Ruan & Douglas, sp. nov. G P. rufiposterus Ruan & Douglas, sp. nov. H P. unguicularis. I P. yunnanensis Ruan & Douglas, sp. nov. J P. zhiweii Ruan, Douglas & Qiu, sp. nov.

Figure 25. Ventral view of pro- and mesothorax of Chinese Phorocardius species. Showing shape of ventral surface of prosternal process (upper, yellow dashed line), mesepisternum (middle, green arrow and dashed line) and projections on posterior edge of mesosternum (lower, red arrow and solid line) middle, green arrows indicating anterior-mesal angle of mesepisternum. A P. alterlineatus Ruan & Douglas, sp. nov. B P. flavistriolatus Ruan & Douglas, sp. nov. C P. florentini. D P. magnus. E P. manuleatus. F P. minutus Ruan & Douglas, sp. nov. G P. rufiposterus Ruan & Douglas, sp. nov. H P. unguicularis. I P. zhiweii Ruan, Douglas & Qiu, sp. nov.

Figure 26. Female abdominal ventrite V in Chinese Phorocardius species (ventral view). A P. alterlineatus Ruan & Douglas, sp. nov. B P. flavistriolatus Ruan & Douglas, sp. nov. C P. florentini. D P. magnus. E P. manuleatus F P. rufiposterus Ruan & Douglas, sp. nov. G P. unguicularis H P. yunnanensis Ruan & Douglas, sp. nov, arrows showing incisions near apex.