Madurasia undulatovittata (Motschulsky) Prathapan, Kaniyarikkal Divakaran, 2016

Prathapan, Kaniyarikkal Divakaran, 2016, Revision of the legume-feeding leaf beetle genus Madurasia Jacoby, including a new species description (Coleoptera, Chrysomelidae, Galerucinae, Galerucini), ZooKeys 597, pp. 57-79 : 63-68

Taxon classification Animalia Coleoptera Chrysomelidae

Madurasia undulatovittata (Motschulsky) comb. n. Figs 1, 3-7, 8-20, 23, 25, 27, 29, 31, 33, 35

Teinodactyla undulatovittata Motschulsky, 1866: 417 [Sri Lanka, Lectotype (ZMUM)]- Wagner and Bieneck 2012: 214-215.

Longitarsus undulatovittatus : Gemminger and Harold 1876: 3509- Maulik 1926: 361.

Monolepta undulattovittata : Ogloblin 1930: 112.

Madurasia obscurella Jacoby, 1886: 381 ["Madura, Madras Presidency", Southern India– Lectotype (BMNH)]- Maulik 1936: 74- Wilcox 1973: 435- Takizawa 1987: 39- Takizawa and Kimoto 1990: 8- Takizawa 1990: 281- Mohamedsaid 1997: 5- Medvedev and Sprecher 1999: 310 (catalogue)- Mohamedsaid 2000: 370- Kimoto 2005: 58- Beenen 2010: 481- Bezděk 2012: 422, 424. New synonym.

Neorudolphia bedfordi Laboissière, 1926: 191 [Brit. Sudan, on Cajanus indicus , Syntype (ZMUH, Hamburg)]- Aslam 1972: 500 (synonymized with Madurasia obscurella Jacoby)- Wilcox 1973: 435- Weidner 1976: 229.

Description.

Body: length 2.0-3.0 mm; width 1.0-1.3 mm; 2.0-2.3 times longer than wide. General color pattern consistent but highly variable in intensity (Figs 1, 3, 5-7). Head dark brown to pale brown, often darker than pronotum. Basal antennomeres 3-6 pale straw brown, distal antennomeres becoming progressively darker. Pronotum more or less pale brown, generally paler than head. Background color of elytron paler than pronotum. Lateral margin of dark elytral stripe emarginate in anterior 1/3 and posterior 1/3; stripe broadening posteriorly, covering width of elytral apex. In some specimens, elytra darker laterally giving the impression of a pale, medially narrowed line on a dark elytron. Intensity of stripe’s darkness varies from pale straw brown (Fig. 1) to dark brown. In type of Madurasia undulatovittata , elytral stripes are hardly visible (Fig. 1). In some specimens, widest region in middle of stripe extends to lateral elytral margin, thus dividing pale colored lateral area into anterior and posterior spots (Figs 6, 7). Ventral aspect (Fig. 4) generally paler than head. Metasternum slightly darker than pro– or mesosternum. Metepisternum darker than metasternum. Abdomen darker laterally and posteriorly in many specimens. In darkest specimens, ventral side dark brown to piceous. Legs pale brown, all femora nearly concolorous with abdominal ventrites; metafemora darker distally in some specimens. All tibiae paler than femora. Metatibia and first metatarsomere whitish in some specimens. Claw tarsomere and bilobed tarsomere often darker than preceding ones.

Antenna (Fig. 9) reaches middle of elytron or a little beyond. Proportionate length of antennomeres 1-11: 1: 0.54-0.57: 0.44-52: 0.65-0.69: 0.59: 0.66: 0.66-0.69: 0.62-0.75: 0.73-0.75: 0.69-0.72: 0.69-0.71: 0.81-0.91. Transverse diameter of eye 5.3-8.8 times width of orbit, 2.9-4.4 times width of interantennal space, 1.7-1.8 times width of antennal socket, 0.6 times distance between eyes. Pronotum (Fig. 14) 1.3 times wider than long, posterior 1.1 times wider than anterior.

Proportionate length of femur–tibia–tarsomeres 1-4 as follows: 1: 1.0-1.1: 0.3: 0.1-0.2: 0.1-0.2: 0.3 (foreleg); 1: 0.9-1.0: 0.3: 0.1-0.2: 0.1-0.2: 0.2-0.3 (midleg); 1: 1.1-1.2: 0.4: 0.1-0.2: 0.1: 0.2 (hindleg). Two visible apical tergites completely exposed in most females, particularly when killed in alcohol.

Posterior margin of apical ventrite in male (Fig. 23) indistinctly lobed medially. Receptacle of spermatheca 1.6 times wider than long (Fig. 29). Tignum not widened proximally (Fig. 33); membranous distal region widest medially.

Aedeagus in lateral view (Fig. 27) strongly curved after basal 1/2, acutely narrowed in proximal 1/3, with weakly curved apex. In ventral view (Fig. 25), widest in proximal 1/3, narrowing sharply towards apex in apical 1/3, lateral margin a little abruptly narrowed preapically. Ventral aspect of aedeagus depressed with a convex portion in middle.

Material examined.

Types. Madurasia undulatovittata : Lectotype ♀. "Teinodactila / undulato / vittata Motch / Ceylon"; " Monolepta / undulatovittata Mots. / 1926 D. Ogloblin det."; "LECTOTYPUS / des Döberl 2005" (ZMUM).

Madurasia obscurella : Lectotype ♀. “Type” (rectangular red label); "Andrewes / Bequest. / B. M. 1922 –221.”; “Madura”,“738” (3 in 738 is not legible as pierced by pin); " Madurasia obscurella Jac. /Type"; “SYNTYPE” (white circular disc with sky blue margin); "Lectotype / Madurasia obscurella Jacoby / des. K. D. Prathapan, 2015" (here designated, specimen on card, right antenna missing) (BMNH).

Paralectotype ♀. "Type / H. T." (white circular disc with red border); “Madura”; "Jacoby Coll. 1909 –28a.”; " Madurasia / obscurella / Jac. Type" (Blue label); “SYNTYPE” (white circular disc with sky blue margin); "Paralectotype / Madurasia obscurella Jacoby / des. K. D. Prathapan, 2015" (BMNH).

Non-type material.

AFRICA: Sudan: ♀ British Sudan, S. R. J. Madani, 22.ix.1923, H. M. Bedford, feeding on ‘adis’ (illegible) sudani leaves / Blue Nile A 3024 / Pres by Imp. Bur. Ent. Brit. Mus. 1925-228 / standing as Neorudolfia (sic) bedfordi ; 1 unsexed R. F. Wadmedanai J. W. Cowland 21/9/32 Shotholing seedlings of Phaseolus mungo / Ent. Coll. C 12147 / AFRICA 250,000 55-G Map / Pres. by Imp. Inst. Ent. BM 1933-415 / Standing as Neorudolphia bedfordi / SUDAN Govt.; 1 unsexed Blue Nile 5429 / Aenk H. H. & D. King 26.5.13 On boot / Pres. by Imp. Bur. Ent. Brit. Mus. 1927-103 / Neorudolphia bedfordi V. Laboissière–Dèt. (all BMNH).

ASIA: Bangladesh: ♀ (India) Dacca, 2.vi.1945, D. Leston; ♀ (India) Dacca, 10.v.1945, D. Leston (both BMNH); India: Andhra Pradesh: 3 unsexed Vizagapatnam Dist., Chipurupalli, B.M. 1924-7; Gujarat: 2♀ Baruch, 10.xii.1987, Pigeon pea, CIE A19617; Navasari, 15.iii.1992, Assoc with cowpea, IIE 22432, Madurasia obscurella Jac det. M. L. Cox 1992 (all BMNH); Karnataka : 1 macerated specimen Belgaum, 1-2.viii. 2008, at light, K. Swamy; 2♀, 1♂ Chikkaballapur, 13°25'48"N, 7°43'12"E 694 mt., 29.viii.2010, Nirmala P., at light (all UASB); Kerala : 7♂, 2 ♀ Vellayani, N 08°25'47.5"E, 76°59'8.3", 21.vii.2015, 18 m, Prathapan KD; 19 ♀, same data except for the date 5.vii. 2015 and ex Green gram (NBAIR, JBC, INPC, BMNH); Maharashtra: 3♀ Bandra, Jayakumar, 1905-152; 2 unsexed Bombay (Mumbai), 79.15; 1♀ Bombay, G. Bryant, 1919-147; 1 unsexed, Poona (Pune), 27.viii.1944, D. Leston, BM 1946-365; 1♀ 21.x.1944, D. Leston, BM 1945-86 (all BMNH); Meghalaya : 1♂, 1 ♀ SW of Cherapunjee, 23°13'15"N / 91°40'E, 500-900 m, 11-12.v.2004, R. Businsky (all JBC); New Delhi: 4♀, 6 unsexed 21.viii.1968, on cowpea, Phaseolus and urd (all BMNH); Rajasthan: 1♀Jodhpur N 26°21'4.6"E, 73°2'39" 5.VIII.2015 255 m, Prathapan K. D. (KAU); 10 unsexed Banswara 24.ix.2015, S. Ramesh Babu (KAU); Uttar Pradesh: Saharanpur Div., Siwalik Hills, 8.iv.1928, H. G. Champion, B.M. 1928-518 (BMNH); Uttarakhand: 1 ♀ Dehra Dun, 8.ix.'16, H. G.Champion, BM. 1953- 156; 1 ♀ Ranikhet, 6-8. '16, H. G.Champion, BM. 1953-156 (both BMNH); West Bengal: 2 unsexed Sarda, H. G. Champion, B.M. 1953-156; 2 unsexed Sunderbans, H. G. Champion, B.M. 1953-156 (all BMNH); Sri Lanka: 2 unsexed, 1♀ Girandurukotte no. 68, 16.xii.86 on cowpea, CIE A18795; 2♀ Maha Illupallama, 1976, R. W. Fellowes, R. W. Fellowes, on Glycine & Vigna , CIE A9047 (all BMNH); Yemen: 2 ♀ Al Hudaydah gov., Jabal Bura Valley forest N. P., (stream valley), 240-350 m, 15°52.4-5'N, 43°24.6-25.2'E, J. Bezdӗk, 4.xi.2010; 1♂, 2♀ Socotra Island, wadi Ayhaft, 12°36.5'N, 53°58.9'E, 200 m, J. Bezdӗk, 7-8.xi.2010 (all JBC).

Distribution.

Africa (Sudan); Asia (Bangladesh, India [Andhra Pradesh, Bihar, Gujarat, Haryana, Karnataka , Kerala , Madhya Pradesh, Maharashtra, Meghalaya , New Delhi, Orissa, Punjab, Rajasthan, Tamil Nadu, Uttar Pradesh, Uttarakhand, West Bengal], Nepal, Sri Lanka, Yemen) (Fig. 35).

Remarks.

Madurasia undulatovittata and Madurasia andamanica sp. n. are very similar. However, they can be separated as follows: eight labral setae present in Madurasia andamanica sp. n. (only six labral setae visible in Madurasia undulatovittata , though eight pores are present); elytral stripes are highly variable in Madurasia undulatovittata , even in specimens from the same locality, collected during the same season and on the same host. The elytral pattern in Madurasia andamanica sp. n. is rather consistent. The stripe in Madurasia undulatovittata is wider apically in specimens where it is well defined, while in Madurasia andamanica sp. n., it is narrowed apically. In Madurasia andamanica sp. n., the stripe is distinct and well defined against the pale background color. Verma (1995) recorded variation in elytral color pattern. Lobe in the middle of the posterior margin of the apical abdominal ventrite in males distinct in Madurasia andamanica sp. n., but poorly distinguishable in Madurasia undulatovittata . The two species can easily be separated by the structure of the aedeagus. In lateral view, the apex of aedeagus of Madurasia andamanica sp. n. is curved and pointed, like the beak of a parrot (Fig. 26), while the same in Madurasia undulatovittata is narrowly rounded, and smoothly curved in apical 1/3 (Fig. 27). The sharply raised ridge on the ventral aspect of the aedeagus in Madurasia andamanica sp. n. (Fig. 24) is characteristic, however, this ridge is absent in Madurasia undulatovittata (Fig. 25). Madurasia andamanica sp. n. is confined to the Andaman Islands and reported to feed on pigeon pea, while Madurasia undulatovittata is transcontinental in distribution and a significant pest of a number of species of pulses in southern Asia and Africa (Sudan).

A photograph of the labels provided by Wagner & Bieneck (Fig. 38a in Wagner and Bieneck 2012) shows three labels, two of which show different information for Madurasia undulatovittata (Fig. 2). Labels currently on the specimen indicate that M. Döberl designated the lectotype in 2005. However, no publication by Döberl could be traced in which this specimen is mentioned. According to Wagner and Bieneck (2012), the lectotype was designated by Wagner, and they provide photographs of both the lectotype and its labels. The photograph (Fig. 38b) in Wagner and Bieneck (2012), confirms that the specimen examined by me is the one designated as lectotype by Wagner (Fig. 1). Moreover, Wagner and Bieneck (2012) also mention that the only other specimen, a paralectotype in Motschulsky’s collection, is a male from which the aedeagus has been dissected and subsequently lost. Dr Wagner’s lectotype designation stands valid as that alone is published ( Wagner and Bieneck 2012). Dr Döberl designated the same specimen as lectotype in 2005 as there was a long gap of nearly a decade between the lectotype designation by Dr Wagner and the publication of the same in Wagner and Bieneck 2012 (T. Wagner and M. Döberl, pers. comm., 2016). The specimen collected by Bedford on 22.ix.1923, identified as Neorudolphia bedfordi by Laboissière from the BMNH, probably belongs to the type series of Neorudolphia bedfordi . The lectotype for Madurasia obscurella is here designated, to have a unique name bearer and standard for its application.

Host plants.

Fabaceae : Cajanus cajan (L.) Millsp. (red gram or pigeon pea); Glycine max (L.) Merr. (soybean); Lablab purpureus (L.) Sweet (= Dolichos lablab L.) ( lablab bean); Vigna aconitifolia (Jacq.) Marechal (moth bean); Vigna mungo (L.) Hepper (= Phaseolus mungo L. = Phaseolus radiatus Roxb. non L.) (black gram); Vigna radiata (L.) R. Wilczek (= Phaseolus aureus Roxb. = Phaseolus radiatus L.) (green gram or moong); Vigna radiata (L.) Wilczek var. sublobata (Roxb.) (= Phaseolus sublobatus Roxb.); Vigna umbellata (Thunb.) Ohwi & Ohashi (rice bean) and Vigna unguiculata (L.) Walp. (= Vigna sinensis (L.) Savi ex Hausskn.) (cowpea).

Biology and management.

Information on the host plants and biology of Madurasia undulatovittata was generated by agricultural entomologists in India, under the name Madurasia obscurella , where it is a widely distributed pest of legume crops across many agro climatic zones. The first record of this species as a pest of pulses is that by Menon and Saxena (1970). According to Naresh and Thakkur (1972), it was reported as a major pest of black gram by Naresh and Nene in 1968. However, there is no mention of this leaf beetle in Naresh and Nene (1968). Saxena et al. (1971) described it as a pest of cowpea, green gram or moong and black gram or urd, indicating that it made holes in the leaf lamina. Other recorded host plants include Glycine ( CAB International Institute of Entomology 1990), moth bean ( Pareek et al. 1983), lablab bean ( Gupta and Singh 1984a, b), pigeon pea ( Saxena 1977, Mishra and Saxena 1983), rice bean ( Satyanarayana et al. 1995a, b) and Vigna radiata (L.) Wilczek var. sublobata (Roxb.) (= Phaseolus sublobatus Roxb.) ( Kalaichelvan and Verma 2005).

Gupta and Singh (1984a, b) provided the first account of its life cycle. They recorded the total life cycle as varying between 32 and 44 days and that it completes two generations a year on green gram. A second, more detailed study of the life history was reported by Oza et al. (1996) on cowpea. Eggs were laid singly on soil near the root zone of the plant. The total duration of the life cycle, from egg to death of adult, varied between 35 and 48 days in males and 43 to 58 days in females.

The growth of plants is retarded by severe foliage injury, especially in young plants ( Srivastava and Singh 1976). Leaf damage on green gram in summer and rainy season crops ranged between 5-10% and 15-50% respectively ( Sinha et al. 1985). Larvae are soil dwelling and feed on root hairs ( Srivastava and Singh 1976; Gupta and Singh 1981). Odak and Thakur (1978) reported larval feeding on the root nodules. Gowda and Kaul (1982) recorded adult feeding on leaves, buds and flowers. Gowda et al. (2006) also observed feeding damage by adults on the buds and flowers of pigeon pea. Reddy and Varma (1986) established transmission of southern bean mosaic virus in cowpea by Madurasia undulatovittata . The success in transmission varied from 25 to 43%.

The extent of damage on black gram, green gram and cowpea was 20-60% ( Srivastava and Singh 1976). This is a common pest of mung bean in the first crop season (kharif) in India, coinciding with the southwest monsoon (June to October) ( Tiwari 1978). Singh and Gupta (1982) estimated damage to the leaves of green gram and black gram. Infestation was more pronounced in black gram than in green gram. Infestation starts when the plants are in the two leaf-stage and the insects remain active until flowering ( Dhuri and Singh 1983, Nayak et al. 2005).

In Haryana, Yadav and Yadav (1983) recorded it from cowpea and Mrig and Singh (1985) observed maximum damage on Dolichos lablab during the third week of September, with the pest disappearing after the first week of November. Feleiro and Singh (1985) carried out yield–infestation studies to fix the critical stages of crops requiring protection. They observed that infestation in summer resulted in heavy yield losses, while the pest attack during the rainy season had no significant effect on yield.

Lal (1985) reviewed information on the biology and control of insect pests of mung bean, including Madurasia undulatovittata , in India.

According to Faleiro et al. (1986), Madurasia undulatovittata is a sporadic, but major pest of cowpea. A peak population of 10.0-10.25 beetles/10 plants in summer and 29.50-30.25 beetles/10 plants in the rainy season were recorded by Gupta and Singh (1993) in green gram. Sahoo and Patnaik (1994) recorded the incidence of insect pests in green and black gram, and their seasonal activity and the extent of damage in Orissa. Madurasia undulatovittata was severe on both the crops in the seedling and vegetative stages, and was the first pest to appear at seedling stage on rice bean, continuing to occur until flowering ( Satyanarayana et al. 1995b). Ganapathy and Durairaj (1995) reported it as an important pest on black gram and green gram in drought prone Pudukottai District, Tamil Nadu. There was more damage in black gram (9.78%) than in green gram (1.45%). However, there are also reports of Madurasia undulatovittata only being a minor pest ( Devesthali and Joshi 1994, Kumar et al. 1998).

Dhuri et al. (1984) observed population buildup of Madurasia undulatovittata under ambient temperature of about 32°C, longer duration of bright sunshine and high relative humidity coupled with intermittent rainfall. Sardana and Verma (1986) showed that maximum temperature and sunshine were negatively, but significantly, correlated with the population of the pest, while rainfall showed a significantly positive correlation. Maximum temperature, minimum temperature, sunshine hours and wind velocity had a significantly negative correlation with damage ( Irulandi and Balasubramanian 1999). Nayak et al. (2004) reported a significantly negative correlation with minimum temperature and relative humidity during population buildup on black gram. The population did not show any correlation with maximum temperature, relative humidity and rainfall, but it was highly and significantly correlated with minimum temperature ( Kumar et al. 2007).

Various cultivars of green gram ( Srivastava et al. 1975, Sahoo et al. 1989, Sahoo and Hota 1991) and black gram ( Sahoo et al. 1989) vary significantly in their susceptibility to the pest. Pandey et al. (1995) reported that varieties with thicker leaves were preferred by the pest.

Chemical control remains the most effective option against this pest. Several broad spectrum insecticides have been tried against Madurasia undulatovittata , with varying degrees of success ( Saxena et al. 1971, Naresh and Thakur 1972, Saxena et al. 1975, Verma and Pant 1975, Verma and Lal 1976, Vyas and Saxena 1977, Verma and Lal 1978, Yadav et al. 1979, Vyas and Saxena 1981, Chaudhary et al. 1981, Rajendran et al. 1981, Vyas and Saxena 1982, Mishra and Saxena 1983, Singh et al. 1983, Singh and Gupta 1984, Sinha 1985, Gattoria and Singh 1988, Rahman 1988, 1991a, b, Chander and Singh 1989, Sinha and Sharma 1989, Verma and Dikshit 1990, Logiswaran and Gopalan 1993, Uddin et al. 1994, Das 1999, Gowda et al. 2006 and Pandey et al. 2007). Application of neem seed kernel extract had no significant effect to increase yield in mung bean ( Yadav et al. 1979). Soundararajan and Chitra (2011) tried biological control agents such as Pseudomonas flourescens and Beauveria bassiana and reported that intercropping black gram with sorghum reduced infestation ( Soundararajan and Chitra 2012).