1 2 3 PINGPING CHEN , NICO NIESER & JEN-ZON HO

1National Museum of Natural History, Naturalis, Leiden, The Netherlands 2Hertog Eduardstraat 16, 4001 RG Tiel, The Netherlands 3Taiwan Endemic Species Research Institute, Taiwan, China

REVIEW OF CHINESE RANATRINAE (: ), WITH DESCRIPTIONS OF FOUR NEW SPECIES OF FABRICIUS

Chen, P.-p., N. Nieser & J.-Z. Ho, 2004. Review of Chinese Ranatrinae (Hemiptera: Nepi- dae), with descriptions of four new species of Ranatra Fabricius. – Tijdschrift voor Entomolo- gie 147: 81-102, figs. 1-47, table 1. [ISSN 0040-7496]. Published 1 June 2004. The Chinese Ranatrinae is reviewed based on specimens examined and literature records. Four new species of Ranatra are described: R. lansburyi sp. n. (SW China and Thailand), R. sterea sp. n. (SW China), R. recta sp. n. (S China) and R. incisa sp. n. (SE China). Diagnoses, distribu- tional and biological notes, a key to Chinese species of Ranatrinae, line illustrations of diagnos- tic features for each species and the dorsal habitus of the different genera are provided. Correspondence: Dr. P.-p. Chen, c/o National Museum of Natural History, Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands. Key words. – Nepomorpha; Cercotmetus; Ranatra; new species; key; China; Thailand.

The family Nepidae, or water scorpions, can be rec- The family Nepidae contains about 250 species, ognized by the following set of characteristics: aquatic and is represented in all zoogeographical regions with bugs with the antennae hidden under the head, the highest species densities in tropical areas. The fam- strongly modified raptorial fore legs, and caudally ily can be divided into two groups. Some authors (e.g. with a single narrow respiratory siphon which cannot Lundblad 1933, De Carlo 1951, 1964) consider these be retracted into the apex of the abdomen. as separate families, namely Nepidae and Ranatridae. There are two main forms of Nepidae. The first one More recent authors (e.g. Lansbury 1972a, 1974, consists of oval or elongate oval, dorsoventrally flat- Schuh & Slater 1995, Polhemus 1995) recognize two tened . They keep their forelegs with a strongly subfamilies: Nepinae (with ten genera) and Ranatri- thickened femur horizontally in front, and combined nae (with four genera). We follow the latter modern with the respiratory siphon indeed remind one of small classifications and treat Ranatrinae as a subfamily of scorpions. Linnaeus (1758) gave the name of Nepa Nepidae. (Latin: scorpion) to the water bugs with prominent The Nepinae contain the real water scorpions, raptorial fore legs. The second form consists of elon- which are flat, more or less oval forms with thickened gate bugs, usually with a subcylindrical body, which front femurs. Three genera of this subfamily have been keep their forelegs with elongated moderately thick- found in China so far. (1) The genus Nepa contains ened femora vertically in front. Fabricius (1790) split five species occurring in the temperate zone of the these subcylindrical forms off from genus Nepa, and Northern Hemisphere, two species occur in the north- gave them the name of Ranatra which is considered to ern part of China. (2) The genus Telmatotrephes Stål, be a fantasy name derived from Rana (Latin: frog). 1854 holds six species, three in South America and Most Ranatrinae species have a subcylindrical body, three in Asia, of which T. chinensis Lansbury, 1972b, hence their vernacular name is water stick insects. the only Chinese species, is only known from the The Ranatrinae occurring in China can be distin- holotype from Fujian Province, SE China. (3) Finally, guished from other taxa of Nepidae by the following the genus Laccotrephes Stål, 1866 is widely distributed diagnosis: with a subcylindrical body, and the width in the Old World tropics and subtropics with about of head across eyes larger than the anterior width of 80 species . This is the dominant genus of Nepinae in pronotum (figs. 1-3). China with seven recorded species, and is distributed

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(De Carlo 1964). In Asia the genus contains 29 species including the species described below. In addi- tion, Australia has three species which are related to various Asian species groups. Judging from its para- mere, R. linearis (Linnaeus, 1758) could be associated with some African species (Poisson 1965). Apart from this species, the Asian species of Ranatra do not seem to be closely related to African or American species. Consequently, speciation probably took place from several centres. As a whole, the Chinese Ranatrinae, with 10 out of 39 species, contains 26% of the described Asian fau- na of this taxon. As stated above, the two species of Cercotmetus are part of an tropical Asian element. Six of the eight species of Ranatra with certainty oc- curring in China fall into four of the species groups defined by Lansbury (1972a). As stated above, R. lin- earis is associated with some African species (Poisson 1965). R. sterea sp. n. does not fit in any of the Asian species groups of Lansbury (1972a) and does not seem to be related to species outside Asia either, thus in our opinion, this species forms a monotypical species group. Aukema & Rieger (1995) included the whole of China into the Palaearctic Region. The Palaearctic Region is usually defined as the part of the Old World roughly north of 25º (in Africa) and 30º (in Asia) northern latitude. The Oriental region contains Sri Lanka and the In- dian Subcontinent, southern China, the Philippines and Indonesia except Irian Jaya (Ellis 1988). In China the Yangtze River is considered to form the demarca- tion between the Oriental and Palaearctic regions (El- lis 1988). According to our knowledge of aquatic bugs, South China (Guangdong, Guangxi), Southeast Fig. 1. Cercotmetus brevipes, , body length 34 mm. China (Fujian, Taiwan), and Southwest China (Guizhou, Sichuan, Yunnan) hold an Oriental fauna. Notably the fauna of Yunnan and Guangxi is related to the fauna of Southeast Asia, especially to the fauna in most areas of the country except for the northeast, of the Mekong basin (Nieser et al. in press). Only west and northwest (Polhemus 1995). Four genera are three species of Ranatra: R. linearis , R. chinensis and R. in the subfamily Ranatrinae, two of them are so far re- unicolor Scott, 1874 have a Palaearctic distribution. stricted to Australia. The genus Cercotmetus Amyot & The latter two extend into the Oriental Region in re- Serville, 1843 is restricted to tropical Asia with one stricted sense. All three Palaearctic species occur in species extending into the subtropics and one sub- China. The total number of species of Ranatra with an species occurring in Australia (Lansbury 1975). Total- Oriental distribution is 29, of which five occur in Chi- ly ten species are described in this genus (Lansbury na: R. falloui Montandon, 1907 extends marginally 1973, Keffer & Polhemus 1999), of which two occur into the eastern Palaearctic; and the four newly de- in China. The name Cercotmetus is derived from the scribed species are presently only known from their Greek substantive ‘kerkos’ (= tail), combined with the type areas in the Oriental part of China, e.g., Guang- Greek verb ‘tmaoo’ (= to cut). It means ‘with a cut tail’ dong, Taiwan and Yunnan. and refers to the respiratory siphon, which is distinct- Apart from some descriptions of species collected in ly shorter than that in the genus Ranatra. China, e.g., R. chinensis and R. falloui , the taxonomic Ranatra is a cosmopolitan genus with about 100 work on Chinese Ranatrinae consists essentially of cat- species. It reaches its highest species density in tropical alogues and the inclusion of most Chinese species in areas, notably in South America with about 45 species the revisions of Cercotmetus and Ranatra (resp. Lans-

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a modern catalogue of Palaearctic Nepidae, citing one species of Cercotmetus and six species of Ranatra as recorded from China. Finally, Hua (2000) published a list of Chinese insects, in which aquatic bugs are treated. Apparently he did not consult the work of Polhemus (1995), resulting in an outdated classifica- tion, including a few spelling and/or printing errors. Keys to the genera of Chinese aquatic Heteroptera can be found in Zheng et al. (1994). However, in their key to the Nepidae, the genus Cercotmetus is lacking. The only keys available to identify Chinese Ranatrinae to species are found in Lansbury (1972a) for Ranatra and Lansbury (1973) for Cercotmetus. However, as Lans- bury (1972a) excluded R. linearis (occurring in the Palaearctic region) from his key to the Oriental species of Ranatra, and we add here four new species for Chi- na, therefore, we present a key and diagnoses of the species actually occurring in China. In addition, diag- noses of two species are presented for further compar- ison, as we strongly doubt their occurence in China.

Biology Two papers dealing with biological notes on Chi- nese Ranatra were published by Hoffmann (1930a, 1930b), in which he reported respectively on R. chi- nensis and R. filiformis Fabricius, 1790. However, judging from his illustration, we consider that his R. filiformis is an misidentification (see under the discus- sion of R. filiformis below). Most Ranatra species live in stagnant waters, such as marshes, ponds and canals, where they usually hide between emergent vegetation awaiting prey. Cloarec (1972) and Cloarec & Joly (1988) studied the behaviour of R. linearis. They found that first instar larvae tend to stay close to the surface film, preferably perching on plants such as Lemna Linnaeus and Elodea Michaux. During devel- Fig. 2. Ranatra longipes, , body length 22.7 mm. opment they prefer gradually deeper sites, and plant preference shifts to plants with distinct stems, such as Phragmites Adanson and Calamagrostis Adanson. Younger larvae prefer green plants, older larvae and bury 1973, 1972a). The catalogue of Palaearctic adults chose brown plants to perch on. The adults Hemiptera by Oshanin (1912) recorded three species wait for their prey with their head down, front legs of Ranatra from China. Esaki (1926) dealt with the stretched forward and the body in line with the stem fauna of Taiwan only, recording three species of Rana- but in an angle of 30˚ or less. When the stored air is tra from the island. Wu (1935) published the first cat- depleted, they walk slowly backwards to the surface alogue of insects specifically for China, and recorded until the respiratory siphon makes contact with the seven species, of which one has been synonymized by air. Gas exchange takes about one minute, after which Lundblad (1933). Hoffmann (1941) published a cata- the walks down again until reaching roughly the logue for China and adjacent areas, citing five species same place as before. Some species occur in streams, of Ranatra for China, omitting R. linearis which was where the bugs stay at the edge of the stream, and included by Wu (1935) based on a record from most often prefer sites between washed out plant Suzhou, eastern China. Lansbury (1972a) reviewed roots. A few species may burrow in mud and catch the Oriental species of Ranatra including all species prey through the surface film (Chen et al. in press.). recorded from China except for R. linearis. Lansbury Ranatra species are carnivorous, maximum prey (1973) revised the genus Cercotmetus and gave the first size increasing with the size of the bug during devel- Chinese record for this genus. Polhemus (1995) gives opment. Minimum prey size is apparently constant,

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first instar larvae feed on small prey, such as water fleas, but adults will still prey on water fleas (Chen et al. in press). R. ‘filiformis’ larvae and adults were re- ported by Hoffmann (1930b) to have a preference for mosquito larvae. He also reported that this species is less sensitive to organic pollution than R. chinensis. Cercotmetus species are usually found in virtually stagnant bays of streams or in pools in stream beds, often among floating plant debris. They are distinctly better swimmers than Ranatra species. Notably larvae constantly move their front legs in front of the body when swimming. This led to the remark by Nieser & Chen (1991) that the fore legs are apparently used for swimming. However, subsequent observations have shown that the propulsion is by the middle and hind legs, the fore legs seem to be used to feel for obstacles in front (N. Nieser personal observation). The eggs of Ranatrinae are characterized by two respiratory horns, whereas eggs of Nepinae have 4-26 horns (Cobben 1968). The eggs of Ranatra are usual- ly inserted in floating plant debris, or soft stems just under the water surface (Poisson 1924, Larsén 1936), but at least R. chinensis is reported to deposit eggs in mud (Hoffmann 1927). Usually the horns point up- ward above the water surface more or less vertically. In Chinese species, this has been confirmed for R. chi- nensis by Hoffmann (1930b) and R. linearis by Larsén (1936). Hoffmann (1930b) noted for his R. filiformis that the eggs are so placed that the horns are parallel with the surface of the water. He also observed that in R. filiformis the hatching nymph lies with its dorsal side upwards, whereas in R. chinensis and R. linearis the hatching nymph has its head upwards. The egg- laying habits of Cercotmetus are unknown.

MATERIAL AND METHODS For this study, the following collections were ex- amined: CCBC P.-p. Chen Collection, Beijing, China. NCTN N. Nieser Collection, Tiel, The Netherlands. NMSC Zoological Reference Collection, National University of Singapore. RMNH National Museum of Natural History, Lei- den, The Netherlands. TESI Taiwan Endemic Species Research Institute, Taiwan, China. ZMAN Zoological Museum, University of Amster- dam, The Netherlands. Fig. 3. R. sterea, holotype, , body length 33.5 mm. Measurements to specimens in this paper are in mil- limeters and presented as a range. In the descriptions of new species, if available, five specimens of each sex were measured. When more than two specimens were measured the mean is added in italics. Specific dimen- sions of the holotype are cited between brackets {}. The body length refers to the length without respira-

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tory siphon, and is measured from the base of the ros- Species list of Chinese Ranatrinae trum to the caudal margin of the seventh tergite in dorsal view. The width of an eye is measured from the Cercotmetus Amyot & Serville, 1843 inner border of the ommatidia to the outer border. C. asiaticus Amyot & Serville, 1843 ...... 86 The length of pronotum and its anterior and posterior C. brevipes Montandon, 1909 ...... 87 lobes are measured along median line in dorsal view. There is a mediodorsally interrupted transverse groove Ranatra Fabricius, 1790 in the posterior half of the prothorax. The anterior R. chinensis Mayr, 1865 ...... 90 and posterior lobes are the parts of the pronotum an- R. falloui Montandon, 1907 ...... 90 teriorly and posteriorly of this groove. The dividing R. incisa sp. n...... 93 line between distal and proximal part of the fore femur R. lansburyi sp. n...... 89 is through the apex of the largest tooth perpendicular R. linearis (Linnaeus, 1758) ...... 90 on the opposite (dorsal) margin. R. recta sp. n...... 97 Although most Ranatrinae are more or less uni- R. sterea sp. n...... 98 formly light to medium brown, some show diagnostic R. unicolor Scott, 1874 ...... 93 colour characteristics. However, very often specimens are totally covered with a thin layer of fine mud which Key to Chinese species of Ranatrinae should be scratched away with a needle or a fine brush 1. Fore femur shorter than length of pronotum; res- in order to check the colour beneath. The colour of piratory siphon ¼ or less than median length of the eyes is castaneous to dark grey and not given in the body in dorsal view, fig. 1, Cercotmetus ...... 2 diagnoses. – Fore femur longer than length of pronotum; res- Specimens are always morphologically macropter- piratory siphon over one third as long as the me- ous. Some species occur in two forms, one with a re- dian length of body in dorsal view, figs. 2, 3, duced prothorax e.g., R. diminuta Montandon, 1907, Ranatra ...... 3 R. occidentalis Lansbury, 1972 and R. varipes Stål, 2. Vertex without tubercle, body length less than 42 1861. It is supposed that the form with a reduced mm, middle femur slightly shorter than length of pronotum has also reduced indirect flight muscles pronotum, female genital operculum not extend- (Lansbury 1972a) but so far differences in the devel- ing beyond apex of abdomen (fig. 43), male para- opment of the flight muscles between such forms have mere fig. 29 ...... C. brevipes not yet been verified. – Vertex with prominent tubercle between eyes, The respiratory siphon is a thin, non retractable tube body length over 45 mm, middle femur about at the caudal end of the body. This tube consists of two 1.5 times length of pronotum, female genital op- halves, which are united in the middle. In living speci- erculum extending distinctly beyond apex of ab- mens the halves are always kept together but, they may domen (fig. 44), male paramere fig. 28 ...... separate in dead, especially in mounted specimens...... C. asiaticus The spines or bristles on the antennae of Ranatra 3. Metasternum broadly and deeply emarginated are variable: thick and stout, long and thin or short posteriorly (figs. 23, 26); fore femur centrally and thin. In addition, the antennae have sensory pits with single tooth (figs. 21, 22) or if femur cen- which are indistinct in some species (figs. 8-13). The trally bidentate then vertex with prominent tu- actual function of both bristles and pits is unclear. bercle between eyes (fig. 4) ...... 4 Usually the bristles on an antenna of a given species – Metasternum not emarginate posteriorly or, if are similar; in some species there may be different narrowly and shallowly emarginated posteriorly types on the same antenna e.g., in R. linearis (fig. 9). (fig. 25) then fore femur centrally bidentate (fig. 17) and vertex between eyes smooth or at most with vestigial tubercle (fig. 5) ...... 5 4. Vertex with prominent tubercle between eyes (fig. 4); respiratory siphon about ⅓ body length Ranatrinae Douglas & Scott, 1865 ...... R. lansburyi The sequence of diagnoses of species actually – Vertex not raised between eyes, without tubercle recorded in China follows Lansbury (1973) for Cer- (fig. 6); respiratory siphon about ⅔ body length cotmetus, and Lansbury (1972a) for Ranatra. Ranatra ...... R. sterea linearis has been placed after R. chinensis. Diagnoses 5. Fore femur with single prominent tooth central- of two species of Ranatra of doubtful occurrence in ly (fig. 14) ...... 6 China have been placed at the end of the diagnoses – Fore femur centrally bidentate (figs. 15, 16, 17).. section...... 7

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4

5

6

Figs. 4-6. Ranatra, head and pronomum in lateral view (scale 1 mm). – 4, R. lansburyi, paratype ; 5, R. incisa, paratype ; 6, R. sterea, holotype .

6. Fore femur with distal tooth which may be small much less (about 0.3) than median length. Api- but distinct (fig. 14). Paramere with shaft cal finger-like projection of paramere narrow and widened in basal part and apical part curved; api- with shallow subapical indentation (fig. 40) ...... cal finger-like part with pointed apex reaching ...... R. falloui basally of well developed subapical tooth (figs. 9. Distance between middle and hind coxae sube- 31, 32). Female genital operculum very long, ex- qual; paramere apically slightly constricted dor- tending well beyond caudal end of connexiva. .... sally and strongly constricted ventrally (fig. 35) ...... R. chinensis ...... R. incisa – Fore femur without distinct distal tooth. Para- – Minimal distance between hind coxae 1.4-1.7 mere with S-shaped shaft; apical finger-like pro- minimal distance between middle coxae; para- jection strongly curved backwards, with small re- mere not constricted dorsally and only slightly curved apical tooth or acute apex not reaching apicoventrally (fig. 36) ...... R. recta basally of the subapical tooth which may be poor- See also under R. filiformis and R. longipes longipes ly developed (fig. 33); female genital operculum extending not or hardly beyond caudal end of Cercotmetus asiaticus Amyot & Serville connexiva ...... R. linearis (figs. 7, 28, 44) 7. Larger tooth of fore femur positioned about half way femur (fig. 15) ...... 8 Cercotmetus asiaticus Amyot & Serville, 1843: 441. – Larger tooth of fore femur positioned clearly Cercotmetus asiaticus; Lansbury 1973: 89 (redescription), Nieser et al. (in press, new record to China). nearer distal end of femur (fig. 17) ...... 9 8. Hind femur not reaching more than half way along sixth sternite; maximum width of metaster- Material examined. – CHINA: Yunnan Prov., Xishuang- num only slightly less (about 0.8) median banna, Mengkuan River, 19.v.2000, leg. H. H. Tan, 1 , 4 larvae (NMSC); Yunnan Prov., Xishuangbanna, Man-e length. Apical finger-like projection of paramere stream, Mengla (21˚4’N, 101˚5’E), 11.vii.2001, leg. H. M. relatively broad and with deep subapical indenta- Jiu, 1 (NMSC); Yunnan, Manchang stream, 63 km from tion (fig. 34) ...... R. unicolor Jinghong (21˚4’N, 101˚5’E) to Menglun (21˚9’N, 101˚2’E), – Hind femur reaching more than 3/4 of way along 30.v.2002, leg. C. M. Yang & P. Chew, 1 (det. C. M. sixth sternite; maximum width of metasternum Yang, NMSC); Yunnan Prov., Xishuangbanna, 67 km from

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Mengla (21˚4’N, 101˚5’E), 1.vi.2002, leg. C. M. Yang & P. and shallowly and narrowly emarginate caudally. Mid- Chew, 2 (det. C. M. Yang, NMSC); Yunnan, Xishuangban- na, Mangle stream, by paddy field, 4.vi.2002, leg. C. M. dle femur 1.5 times the length of pronotum. Yang & L. Gao, 3 larvae (det. C. M. Yang NMSC). Male. – Paramere as in fig. 28. Female. – Genital operculum extending distinctly beyond the apex of abdomen (fig. 44). Diagnosis Body length 44-53 mm, length of respiratory Distribution siphon 9-17 mm. Colour more or less uniformly pale China: Yunnan, Malaysia, Thailand and the Sun- to medium brown. Fore femur and tibia usually band- daland part of Indonesia. Judging from collections in ed with darker brown, distinct in most specimens. several museums, this species seems to be especially Middle and hind femur and tibia less often and less common in Java (Indonesia), which is also the type distinctly banded with darker brown than fore leg. locality. Vertex with a prominent tubercle between eyes; inte- rocular space 1.1-1.2 times the width of an eye; second Cercotmetus brevipes brevipes Montandon antennal segment without a fingerlike process (fig. 7). (figs. 1, 29, 43) Transverse groove on pronotum short and shallow. Length of pronotum twice or slightly less than twice Cercotmetus brevipes Montandon, 1909: 65. the length of fore coxa and 1.3-1.4 times the length of Cercotmetus brevipes; Lansbury 1973: 92 (redescription, syn- onymy); Polhemus, 1995: 17 (catalogue). fore femur. Fore femur with a single tooth which is Cercotmetus brevipes brevipes; Lansbury 1975: 17. two fifth or slightly more of fore femoral length re- moved from distal apex of femur. Prosternum with a Material examined. – CHINA: Guangdong Prov., Taishan median and two lateral keels, ventrally densely beset County, Shuibu Town (22˚3’N, 112˚7’E), Shuijikeng, with small nodules, and becoming obsolete posterior- 17.vi.1993, leg. P.-p. Chen & Z.-q. Zhu, C9313, 1, 1, 6 ly; in addition, there is a paired row of nodules lateral- larvae (CCBC). Guangdong Prov., Guangzhou (23˚1’N, ly on prothorax, also becoming obsolete posteriorly. 113˚2’E), Botanical Garden of South China, pond, 3. viii.1990, leg. P.-p. Chen, C9004, 1 larva (CCBC). Metasternum with a distinct median longitudinal keel Diagnosis Body length 32-41 mm; length of respiratory siphon 6.5-8.9 mm (fig. 1). Colour more or less uniformly Table 1. Mean value of length of leg segments of Ranatra pale to medium brown. Fore femur and tibia of all legs species described in this publication. n = 2 exx., * n = 3 exx., banded with darker brown, in middle and hind leg ** n = 5 exx. sometimes indistinctly. Vertex between eyes smooth; coxa femur tibia tarsus claw interocular space 1.4-1.7 times the width of an eye: sec- ond antennal segment with a short but distinct finger- R. lansburyi like projection. Transverse groove on pronotum short fore leg * 5.2 8.3 3.4 0.63 and shallow. Length of pronotum more than twice the middle leg * 13.2 12.0 1.93 0.57 length of fore coxa and 1.5 times the length of fore fe- hindleg * 13.5 14.8 2.10 0.71 fore leg 5.9 8.8 3.7 0.66 mur. Fore femur with a single tooth halfway between middle leg 14.8 13.7 2.18 0.66 distal and proximal apex of fore femur. Prosternum hind leg 15.2 17.0 2.33 0.67 with a median and two lateral keels, the lateral keels R. incisa** ventrally sparsely beset with small nodules and, espe- fore leg 5.4 8.6 3.2 0.67 cially the median keel, becoming obsolete posteriorly; middle leg 11.1 12.9 1.92 0.55 prothorax laterally smooth or with indication of a keel hind leg 11.7 13.5 1.87 0.62 fore leg 6.0 9.3 3.3 0.67 anteriorly and posteriorly. Metasternum with a distinct middle leg 12.8 14.8 2.01 0.59 median longitudinal keel which is emarginate shallow- hind leg 13.1 14.7 1.90 0.60 ly and narrowly at caudal margin. Middle femur 0.8- R. recta 0.9 times the length of pronotum. fore leg 5.2 8.0 2.9 0.63 Male. – Paramere as in fig. 29. middle leg 11.0 12.3 1.84 0.57 Female. – Genital operculum not extending be- hind leg 11.7 13.5 1.80 0.66 fore leg 5.5 8.2 3.1 0.65 yond the apex of abdomen (fig. 43). middle leg 11.4 12.7 1.88 0.60 hind leg 10.7 14.0 1.90 0.69 Distribution R. sterea China: Fujian and Taiwan (Lansbury 1973), fore leg 6.7 11.6 4.7 0.90 Guangdong. From Northeast India through conti- middle leg 11.4 10.0 2.05 0.90 nental Southeast Asia to Sumatra, Java, Borneo and hind leg 12.0 13.7 2.30 0.98 the Philippines.

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7 89

10

11 12 13

Figs. 7-13. Ranatrinae, dorsal view of right antenna (scale 0.1 mm). – 7, Cercotmetus asiaticus ; 8, Ranatra lansburyi, paratype ; 9, R. linearis ; 10, R. recta, paratype ; 11, R. sterea, paratype ; 12, R. incisa, paratype ; 13, R. filiformis .

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Ranatra lansburyi sp. n. shorter than pronotum in females (see dimensions), (figs. 4, 8, 18, 23, 30, 41, 45) with a single, moderately developed tooth, approxi- mately in apical two fifth (fig. 18). Width of fore fe- Type material. – Holotype (RMNH) macropterous mur measured from dorsal margin to apex of tooth male, THAILAND: Chiang Mai Prov., Chiang Dao subequal to widest part of femur proximally. Dist., Ban Yang Thung Pong, 8.xi.1995, leg. N. Male. – Hind femur reaching the basal quarter to Nieser, N9508. – Paratypes, same data as holotype, half of genital operculum. Operculum in ventral view 1 2 (NCTN). – CHINA: Yunnan, Xishuangbanna, elongate with a median keel which is distinct in cau- Mengla 21˚4’N, 101˚5’E), stream QD2, leg. H. M dal half; in lateral view boat-shaped with a small Jiu, XTBG, 1 (NMSC). slightly upturned knob caudally, just reaching the Type locality. – Chiang Mai Province, Thailand. base of respiratory siphon. Genital capsule see fig. 41; paramere with apical part deeply incised, resulting in Description two teeth (fig. 30). Dimensions. – Body length 27.5-29.2-30.4 mm Female. – Hind femur just not reaching the genital {29.8}, 31.6-33.6 mm; respiratory siphon 10.4- operculum. Operculum distinctly projecting beyond 11.5-12.7 mm{12.7}, 12.0-12.5 mm; width of head apex of abdomen (fig. 45). 2.53-2.65-2.86 mm {2.65}, 2.80-2.81 mm; width of an eye 0.73-0.75-0.80 mm, 0.80-0.80 Biological notes mm; interocular space 1.05-1.13-1.26 mm, 1.20- The habitat of the holotype is a mountain stream, 1.22 mm; anterior width of pronotum 2.10-2.18- width 10-15 m, depth mostly up to 0.5 m, the speci- 2.32 mm 2.32-2.33 mm; humeral width of prono- mens were collected from washed out plant roots and tum 2.68-2.70-2.73 mm {2.70}, 3.12-3.20 mm; entangled plant debris at the edges of the stream. length of pronotum 7.6-8.0-8.3 mm {8.3}, 9.0- 9.1 mm; length of anterior lobe of pronotum 4.9- Distribution 5.3-5.6 mm, 6.0-6.2 mm; length of posterior lobe Southwest China (Yunnan) and Northern Thai- 2.6-2.7-2.7 mm, 2.8-3.1 mm. Fore leg: mean val- land (Chiang Mai). ue of length of distal part of femur 3.6 mm, 4.0 mm; proximal part of femur 4.7 mm, 4.9 mm. Comparative notes Dimensions of leg segments are given in table 1. This species belongs to the R. gracilis-group of Lans- Structural characteristics. – Clypeus higher than lo- bury (1972a), characterized by the tuberculate vertex rum, clypeoloral sulcus distinct and deep; vertex with and broadly emarginate posterior margin of the a distinct tubercle between eyes (fig. 4). Eye distinctly metasternum. R. lansburyi is most similar to R. gracilis narrower than interocular space. Antenna as in fig. 8. Dallas, 1850 and R. parmata Mayr, 1865. Both Anterior margin of pronotum (‘collar’) raised in the species have the subapical part of the paramere of middle, anterior lobe of pronotum about twice as long semicircular shape, not incised, although the paramere as posterior lobe, sublateral swellings caudally on of R. parmata has a small tooth ventrally on the sub- pronotum indistinct to virtually absent. Scutellum apical part (fig. 39). Females of R. gracilis are smaller twice as long as broad, dorsal surface smooth with a than those of R. lansburyi with a shorter siphon (29 pair of depressions in caudal half. Prosternum with a and 8.5 respectively) and the mid-tibia is slightly broad shallow longitudinal trough and a rather low longer than the mid-femur in R. gracilis. Males of R. and blunt but distinct (except in the male paratype gracilis, apart from the difference in parameres and the from Thailand) keel which ends close to the conspicu- ratio mid-tibia to mid-femur mentioned above, have ous prothoracic pit. Mesosternum with a pair of blunt the hind femur not surpassing the end of the mem- nodules at its anterior margin, caudal half with a shal- brane as in R. lansburyi. R. parmata and R. lansburyi low median fovea, widening posteriorly. Metasternum are very similar, the only reliable character is the dif- with a pair of low, submedian, posteriorly converging, ference in male parameres (figs. 30, 39). In addition, carinae in its anterior two thirds; posterior margin dis- the hind femur of R. parmata reaches only about tinctly emarginate (fig. 23). Smallest distance between halfway the sixth sternite in both sexes, whereas in R. middle coxae divided by the smallest distance between lansburyi it reaches near the posterior margin of the hind coxae is 2 in males (0.95/0.47 mm) and 1.5 sixth sternite in females and onto the genital opercu- (1.2/0.8 mm) in females. Membrane reaching to or lum in males. just beyond posterior margin of sixth tergite (fig. 45). Fore femur annulated light and dark brown, remain- Etymology der of legs apparently unicolorous (the legs covered This species is named in honour of Dr. Ivor Lans- with a thin but compact layer of dried mud). Fore fe- bury (Oxford, U.K), for his outstanding contribution mur slightly longer than pronotum in males, slightly to the knowledge of aquatic Hemiptera.

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17

18

19

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21 22

Figs. 14-22. Ranatra, fore femur, tibia and tarsus (scale 1 mm). left, external view (14, 15, 17, 18, 19, 21), left, internal view (20), right, internal view (16), detail of left fore femur (22). – 14, R. chinensis ; 15-16, R. unicolor ; 17, R. incisa, holotype ; 18. R. lansburyi, paratype ; 19-20, R. filiformis, ; 21-22, R. sterea, paratype .

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Ranatra chinensis Mayr Ranatra linearis (Linnaeus) (figs. 14, 31, 32) (figs. 9, 27, 33) Ranatra chinensis Mayr, 1865: 446. Nepa linearis Linnaeus, 1758: 441. Ranatra chinensis; Hoffmann 1941; 55 (distribution in Chi- Ranatra linearis; Fabricius 1790: 2; Kanyukova 1989: 121 na); Lansbury 1972a: 301 (key, redescription, syn- (key, distribution in former USSR); Polhemus 1995: 18 onymy); Kanyukova 1989: 124 (distribution in Russia); (general distribution). Polhemus, 1995: 17 (general distribution).

Material examined. – CHINA: Heilongjiang Prov., small Material examined. – China: Xinjiang Autonomous Re- gion, Buerjin (47˚7’N, 86˚9’E), NW of Buerjin Town, ponds in the valley of the Mohe River, 1-10.viii.1939, leg. M. Nikitin, 1, det. I. Lansbury (NCTN); Hubei Prov., small pond near bridge, 21.viii.1991, leg. P. Chen, 2 1 Chang-yang County (30˚4’N, 111˚1’E), Jin-yang-kou (NCTN). Town, fish pond, 24.viii.1990, leg. Chuan-Ren Li, 2, 1 (CCBC); Sichuan Prov., Leshan (29˚6’N, 103˚7’E), Wuyou- ba, canal near Daduhe hotel, 29.vii.1989, turbid water, sam- Diagnosis ple between submerged terrestrial vegetation, leg. P. Chen & Body length 30-41 mm, length of respiratory N. Nieser, N8958, 1 (NCTN); Sichuan Prov., Leshan, siphon 30-36 mm. Colour more or less uniformly pale 31.vii.1989, overflown edges of stream with sparse grasses, brown, prosternum pale reddish to reddish brown. leg. P. Chen & N. Nieser, N8961, 1 (NCTN); Sichuan Legs not annulated except for distal dark brown to Prov., Jiu Zhaigou (33˚2’N, 103˚9’E), alt. 1400-2000 feet, blackish rings on segments of middle and hind legs. viii-ix. 1929, leg. D. G. Graham, 1, det. I. Lansbury (NCTN); Yunnan Prov., Lijiang (26˚8’N, 100˚2’E), N8970, 2 Vertex without tubercle between eyes; eye width dis- larvae (NCTN); Yunnan Prov., 10 km NE Kunming (25˚0’N, tinctly less than interocular space. Antenna as in fig. 9. 102˚7’E), Jindian water reservoir, ca. 200x750m, edges Transverse groove on prothorax inconspicuous; flooded grassland, 27 viii. 1989, leg. P. Chen & N. Nieser, pronotum smooth; length of pronotum 1.1-1.2 times N8974, 3 (NCTN); JAPAN: Tsushima, September-October, the length of fore coxa. Length of fore coxa about two H. Frühstorfer, 1 (ZMAN). thirds the length of fore femur; fore femur with a sin- gle tooth which is about two fifth of fore femoral Diagnosis length removed from distal apex of femur. Proster- Body length 40-53 mm, length of respiratory num with a median longitudinal carina. Metasternum siphon 35-57 mm. Colour more or less uniformly pale distinctly elevated, not emarginate caudally. brown, in some specimens the anterior lobe of the Male. – Paramere as in fig. 33. prothorax is somewhat darker than posterior lobe. Female. – Genital operculum (fig. 27) not extend- Legs not annulated except for distal dark brown to ing beyond the apex of abdomen. blackish rings on segments of middle and hind legs. Vertex without tubercle between eyes; eye width sube- Distribution qual to interocular space. Transverse groove on pro- China: Jiangsu (Wu 1935) and Xinjiang. Probably thorax shallow; pronotum with a faint median keel on widespread in the western part of the Chinese Palearc- anterior lobe, extending onto posterior lobe; length of tic area. Roughly south of 60˚N in Europe, North pronotum 1.1-1.2 times the length of fore coxa. Africa, Caucasus, Asia Minor, Central Asia, Russia Length of fore coxa about two thirds the length of fore south of 60˚N and west of 92˚E (Kanyukova 1989, femur; fore femur with a single large tooth which is Polhemus 1995). slightly less than two fifth of fore femoral length re- moved from distal apex of femur; in addition, there is Ranatra falloui Montandon a small subapical tooth (fig. 14). Prosternum with a (fig. 40) median longitudinal keel. Metasternum distinctly ele- vated, centrally depressed, not emarginate caudally. Ranatra falloui Montandon, 1907: 54. Male. – Paramere as in figs. 31-32. Ranatra falloui; Hoffmann 1941: 56 (distribution in Chi- na); Lansbury 1972a: 314 (redescription, synonymy); Female. – Genital operculum extending distinctly Polhemus, 1995: 17 (general distribution). beyond the apex of abdomen. Material examined. – CHINA: Hunan Prov., Changsha Distribution (28˚2’N,112˚9’E), Agricultural School of Changsha, pond, China: distributed throughout China except 24.ix.1991, leg. P. Chen, C9112, 1 (NCTN); Hubei Prov., Wuchang Dist., Foutou Lake, 30˚3’N, 114˚3’E, 16.x.1991, Northwest China (Gansu, Ningxia, Qinghai, Xin- jiang) and Xizang (Tibet). Japan, Korea, Russia (Far leg. P. Chen, C9117,1 (NCTN). East), Burma and India (Polhemus 1995). Diagnosis Body length 27-34 mm, length of respiratory siphon 17-23 mm. Colour more or less uniformly pale

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23 24

25

26 27

Fig. 23-26. Ranatra, meso- and metasternum (scale 1 mm). – 23, R. lansburyi, paratype ; 24, R. incisa, holotype ; 25, R. recta, paratype ; 26, R. sterea, paratype . – Fig. 27. Ranatra linearis, , ventral view of genital capsule.

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brown, except for anterior lobe of pronotum usually Diagnosis somewhat darker than posterior lobe. Legs not annu- Body length 23-34 mm, length of respiratory lated except for distal dark brown to blackish rings on siphon 16-26 mm. Colour variable from pale to dark segments of middle and hind legs. Vertex dorsally be- brown. Most specimens more or less uniformly pale tween eyes smooth; eye width slightly less than inte- brown, scutellum usually darker greyish brown. Legs rocular space. Transverse groove on prothorax shallow not annulated except for distal dark brown to black- but well visible; anterior lobe of pronotum sublateral- ish rings on segments of middle and hind legs; fore fe- ly with a pair of shallow longitudinal impressions mur varying from uniformly pale brown to strongly which are converging posteriorly; length of pronotum mottled with medium brown, notably on the median 1.05-1.11 times the length of fore coxa. surface. Vertex in some specimens somewhat pointed Length of fore coxa slightly less than two thirds the dorsally between eyes but without distinct tubercle; length of fore femur; fore femur with two large teeth eye width slightly less than interocular space. Trans- which are slightly more than two fifth of fore femoral verse groove on prothorax distinct and deep; anterior length removed from distal apex of femur, the smaller lobe of pronotum sublaterally with a pair of longitu- tooth visible in median view (as in R. unicolor, figs. 15- dinal impressions which are converging posteriorly; 16). Hind femur in males reaching or nearly reaching length of pronotum 1.02-1.05 times the length of the posterior margin of sixth sternite, in females reach- fore coxa. Length of fore coxa slightly less than two ing over halfway sixth sternite. Prosternum with a thirds the length of fore femur; fore femur with two weak median longitudinal keel in anterior half. large teeth which are slightly more than two fifth of Metasternum not elevated, smooth, not emarginate fore femoral length removed from distal apex of fe- caudally. mur, the smaller tooth visible in median view (figs. Male. – Paramere as in fig. 40. 15, 16). Hind femur in males nearly reaching halfway Female. – Genital operculum not extending be- sixth sternite, in females somewhat shorter. Proster- yond the apex of abdomen. num with a weak median longitudinal keel in anteri- or half. Metasternum not elevated, smooth, not Remarks emarginate caudally. This species is very similar to R. unicolor, see there. Male. – Paramere as in fig. 34. Female. – Genital operculum not extending be- Distribution yond the apex of abdomen. China: Jiangsu, Hubei, Hunan, Fujian, Taiwan, Sichuan, Xizang (= Tibet). Vietnam, Thailand. Remarks This species is very similar to R. falloui. The most reliable difference is in the parameres (figs. 34, 40). In Ranatra unicolor Scott addition, the hind femur is in R. falloui somewhat (figs. 15, 16, 34) longer than in R. unicolor. Ranatra unicolor Scott, 1874: 452. Ranatra unicolor; Hoffmann 1941: 58 (distribution in Chi- Distribution na); Lansbury 1972a: 312 (redescription, synonymy); Polhemus 1995: 18 (general distribution). China: distributed throughout China with the pos- sible exception of Qinghai and Xizang. Japan, Korea, Russia (Far East and the extreme Southeast of Euro- Material examined. – CHINA: Tianjin (39˚1’N, 117˚2’E), pean Russia), Central Asia, Caucasus, Saudi Arabia, ditch at SW of the city, between reeds, 14.vii.1989, leg. N. Iran, Iraq (Kanyukova 1989, Polhemus 1995). Nieser, N8950, 1 1 (NCTN); Gansu Prov., Wen Xian (33˚0’N, 104˚6’E), vii.1988, 1 (CCBC). Sichuan Prov., Le- shan (29˚6’N, 103˚7’E), small stream draining a rice field, Ranatra incisa sp. n. width 0.5 m depth 0.2-1 m, between emergent vegetation, (figs. 5, 12, 17, 24, 35, 42, 46) 30.vii.1989, leg. P. Chen & N. Nieser, N8960, 1 (NCTN); Sichuan Prov., Leshan, overflown banks of stream, between Type material. – Holotype (RMNH), macropterous sparse grasses, 31.vii.1989, leg. P. Chen & N. Nieser, male, CHINA: Taiwan Prov., Pingtung Co. (22˚6’N, N8961, 1 (NCTN); Yunnan Prov., Lijiang (26˚8’N, 102˚5’E), Neipu, 20.ix.2003, leg. J.-R. Wu. – 100˚2’E), 1 km along the road to Dali, ditch with slow cur- rent along the road, rich aquatic and emergent vegetation, Paratypes: same data as holotype 14 15 (RMNH, bottom clay with plant debris, 16.viii.1989, leg. P. Chen & CCBC, NCTN, TESI); Taiwan Prov., Pingtung Co., N. Nieser, N8968, 1 (NCTN); Yunnan Prov., 10 km NE Neipu, 1.vii.2003, leg. J.-R. Wu , 1, 1; Taiwan, Kunming (25˚0’N, 102˚7’E), Jindian water reservoir, ca. Yilan Co.(24˚7’N, 121˚7’E), Shuanglianpi, 23.ii. 200x750m, edges flooded grassland, 27 viii. 1989, leg. P. 1985, leg. T.-J. Chen, 1 3 (CCBC, NCTN). Chen & N. Nieser, N8974, 1 (NCTN). Type locality. – Taiwan Province of China.

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29

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Figs. 28-30. Ranatrinae, left paramere, external view (scale 0.5 mm). – 28, Cercotmetus asiaticus; 29, C. brevipes; 30, Ranatra lansburyi.

Description about 1.5 times as long as posterior lobe; humeral Dimensions. – Body length 21.4-21.9-22.2 mm width of pronotum about 1.25 times anterior width in {22.2}, 22.7-24.9-25.9 mm; respiratory siphon males, 1.4-1.5 times in females. Scutellum twice as 15.2-16.8-18.2 mm {18.2}, 17.1-18.7-20.1 mm; long as broad, dorsal surface smooth with a pair of width of head 2.28-2.32-2.35 mm {2.34}, 2.42- shallow depressions in caudal half. Prosternum with a 2.54-2.62 mm; width of an eye 0.76-0.78-0.80 broad shallow longitudinal trough in anterior quarter mm, 0.80-0.85-0.88 mm; interocular space and a rather low and blunt median keel which is dis- 0.77-0.78-0.78 mm, 0.82-0.85-0.88 mm; anterior tinct in anterior half; prothoracic pit rather small but width of pronotum 1.47-1.48-1.50 mm, 1.51- distinct. Mesosternum, anterior margin without or 1.62-1.70 mm; humeral width of pronotum 2.05- with a pair of very weak nodules; caudal half with a 2.07-2.10 mm {2.07}, 2.11-2.32-2.50 mm; length shallow oval to elongate median depression in front of of pronotum 5.3-5.4-5.5 mm {5.5}, 5.2-5.6-5.9 mesocoxae, continuing as a groove between the coxae, mm; length of anterior lobe of pronotum 3.3-3.4- becoming shallow and indistinct anteriorly on 3.5 mm {3.5}, 3.3-3.6-3.8 mm; length of posterior metasternum; posterior margin virtually straight. lobe 2.0-2.0 mm, 1.9-2.1-2.3 mm. Fore leg: Metasternum caudally produced between hind coxae, mean value of length of distal part of femur 3.6 anteriorly with a shallow elongate median depression, mm, 3.8 mm; proximal part of femur 5.1 mm, posteriorly with a broad median keel, posterior margin 5.5 mm. Dimensions of leg segments see table 1. truncate, straight (fig. 24) in half of the specimens, Structural characteristics. – Clypeus higher than lo- with a small somewhat variable notch in other speci- rum, clypeoloral sulcus distinct anteriorly, becoming mens. Smallest distance between middle and hind shallow and indistinct posteriorly. Vertex raised above coxae subequal, about 0.25 mm. Membrane reaching eyes, in one male and two females with a vestigial tu- just beyond posterior margin of sixth tergite (fig. 46). bercle between eyes (fig. 5). Width of eye subequal to Legs unicolorous yellowish brown with distal apices of interocular space. Antenna as in fig. 12. Pronotum, segments darker except for fore tibia. Fore femur (fig. anterior margin (‘colllar’) raised in the middle; anteri- 17) centrally bidentate, slightly over 1.5 times the or lobe with a vestigial median keel extending from length of pronotum, larger tooth approximately in collar to transverse groove; anterior lobe of pronotum apical two fifth. Width of fore femur measured from

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dorsal margin to apex of larger tooth slightly narrower ences in parameres (figs. 35, 36, 38). In addition, R. than widest part of femur proximally. Middle and longipes from Java (the type locality) are slightly larger: hind tibiae longer than their respective femurs (see di- length 22.5-23.9 mm, 26.0 mm; length of mensions); hind tibia reaching halfway to nearly the siphon 19.7-20.9 mm 25.3 mm. However, mea- caudal tip of genital operculum in both sexes. surements by the first two authors show that other Male. – Operculum in ventral view elongate with a populations of this widespread species sometimes con- low and blunt median keel over its entire length; in sist of smaller specimens. The paramere of R. natu- lateral view boat-shaped with a small slightly upturned naensis Lansbury, 1972, although its distal finger-like knob caudally, just reaching the base of respiratory projection is slightly expanded apically, is quite similar siphon. Genital capsule see fig. 42; paramere (fig. 35) to that of R. incisa. R. natunaensis is slightly larger than with distal finger-like projection not expanded apical- R. incisa (body length 23-27 mm, 27-29 mm) ly; dorsal margin of paramere distally with a shallow and can be distinguished by the distance between the concavity, paramere ventrally incised just proximally middle coxae being much smaller (‘almost contigu- of the curved head of paramere resulting in a small ous’, Lansbury 1972a), than the distance between the tooth with a rectangular apex. hind coxae, whereas in R. incisa these distances are Female. – Operculum laterally compressed, with a subequal. Apart from specimens identified as R. sharp and distinct ventral keel and sharp caudal tip longipes longipes and R. recta the geographically most reaching the base of respiratory siphon, not project- close taxon is R. longips thai Lansbury (1972a). Para- ing beyond apex of abdomen (fig. 46). meres of this species are widest and smoothly rounded just proximally of its curved tip whereas as in R. incisa Comparative notes and to a lesser extent R. recta the paramere is narrowed See also under R. recta sp. n. This species belongs to subapically with a small blunt tooth. the R. biroi-group of Lansbury (1972a). It is most closely related to R. recta sp. n. and R. longipes Stål Distribution (1861). The only reliable characteristics are the differ- China: probably endemic to Taiwan.

31

32

Figs. 31-32. Ranatra chinensis, left paramere (scale 0.5 mm). 31, external view; 32, internal view.

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Figs. 33-38. Ranatra, left parameres, external view (scale 0.5 mm). – 33, R. linearis; 34, R. unicolor; 35, R. incisa, paratype; 36, R. recta, holotype; 37, R. filiformis; 38, R. longipes.

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Etymology with a wide, shallow median groove in front of meso- Incisa (Latin adjective, feminine form of incisus, coxae, continuing as a groove between the coxae and meaning incised) refers to the subapical incision of extending onto the anterior part of the metasternum; the paramere. posterior margin virtually straight. Metasternum cau- dally produced between hind coxae, anteriorly with a shallow elongate median depression, posteriorly with a Ranatra recta sp. n. broad median keel, posterior margin narrowly trun- (figs. 10, 25, 36) cate, straight or with a minute median notch (fig. 25). ? = Ranatra filiformis sensu Hoffmann 1941: 56. Smallest distance between hind coxae 1.4 times the distance between middle coxae (0.26/0.19 mm). Type material. – Holotype (RMNH), macropterous Membrane reaching just beyond posterior margin of male (tarsi of hindlegs lacking), CHINA: Guangdong sixth tergite. Legs unicolorous yellowish brown with Prov., Taishan Co., Shuibu Town (22˚3’N, 112˚7’E), distal apices of segments of middle and hind legs Shuijikeng, 17. viI.1993, leg. P. Chen & Z.-q. Zhu. slightly darker. Fore femur centrally bidentate, slight- C9313. – Paratypes: Guangdong Prov., Guangzhou ly over 1.5 times the length of pronotum, larger tooth (23˚1’N, 113˚2’E), Botanical Garden of South China, approximately in apical two fifth. Width of fore femur pond, 3.viii.1990, leg. P. Chen, C9005, 1 2, all measured from dorsal margin to apex of larger tooth teneral (CCBC, NCTN). slightly narrower than widest part of femur proximal- Type locality. – Guangdong Province of China. ly. Middle and hind tibiae longer than their respective femurs; hind tibia reaching halfway to nearly the cau- Description dal tip of genital operculum in both sexes. The male paratype is a strongly shrivelled dry Male. – Operculum in ventral view elongate with a mounted teneral specimen, several body dimensions low and blunt median keel over its entire length; in could not reliably be measured. If no range is given for lateral view boat-shaped with an indistinct, slightly a dimension of males it refers to the holotype. Other- upturned knob caudally, just reaching the base of res- wise the larger dimension refers to the holotype. piratory siphon. Paramere (fig. 36) with distal finger- Dimensions. – Body length 22.8 mm, 21.8- like projection not expanded apically; dorsal margin 23.2 mm; respiratory siphon 16.9-18.0 mm, of paramere virtually straight, paramere ventrally with 14.0-16.7 mm; width of head 2.26-2.33 mm a shallow concavity just proximally of the curved head 2.42-2.46 mm; width of an eye 0.78-0.79 mm, of paramere resulting in a small tooth with a blunt 0.83-0.86 mm; interocular space 0.70-0.75 mm, apex. 0.76-0.82 mm; anterior width of pronotum 1.51 mm, Female. – Operculum laterally compressed, with a humeral width of pronotum 2.22 mm, length of sharp and distinct ventral keel and sharp caudal tip pronotum 5.3 mm, length of anterior lobe of prono- reaching the base of respiratory siphon, not project- tum 3.35 mm, length of posterior lobe 1.95 mm. Fore ing beyond apex of abdomen. leg: mean value of length of distal part of femur 3.3 mm, 3.4 mm; proximal part of femur 4.8 mm, Comparative notes 4.9 mm. Dimensions of leg segments see table 1. According to Lansbury (1972a), this species should Structural characteristics. – Clypeus higher than lo- fall into the R. biroi-group. In our opinion, several sub- rum, clypeoloral sulcus distinct anteriorly, becoming species of R. longipes form a species complex, which will shallow and indistinct posteriorly. Vertex raised above be discussed in a forthcoming publication by the au- eyes with a vestigial tubercle between eyes (virtually thors. The R. longipes-complex consists of a number of identical with R. incisa, fig. 5). Width of eye slightly species which are only separable by the parameres. larger than interocular space. Antenna as in fig. 10. Within it R. recta is most similar to R. incisa and R. Pronotum, anterior margin (‘collar’) raised in the mid- longipes. The only reliable characteristics are the differ- dle; anterior lobe with a vestigial median keel extend- ences in parameres (figs. 35, 36, 38). In addition, the ing from collar to transverse groove; anterior lobe of distance between the middle and hind coxae is sube- pronotum 1.7 times as long as posterior lobe; humer- qual in R. incisa and R. longipes, whereas in R. recta the al width of pronotum 1.5 times anterior width. distance between the hind coxae is 1.4 times that be- Scutellum 1.75 times as long as broad, dorsal surface tween the middle coxae. R. recta is slightly larger and smooth with a pair of shallow depressions in caudal broader than R. incisa. R. natunaensis has the distance half. Prosternum with a broad shallow longitudinal between middle coxae also narrower than between groove trough in anterior quarter, and a rather low hind coxae, but in this species the paramere is ventrally and blunt median keel which is distinct in anterior distincty narrowed, similar as in R. incisa. For compar- half; prothoracic pit shallow. Mesosternum, anterior ison with R. longipes thai see under R. incisa. margin with a pair of very weak nodules; caudal half

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39

40 41

42

Fig. 39. Ranatra parmata, left paramere, external view (scale 0.5 mm). Fig. 40. R. falloui, apex of left paramere (adapted from Lansbury, 1972, fig. 121). Figs. 41-42. Ranatra, male genital capsules, left paramere removed (scale 1 mm). – 41. R. lansburyi, paratype; 42, R. incisa paratype.

Distribution Description China: Guangdong, ?Fujian (see discussion under Dimensions (holotype first, females only). – body R. filiformis). length 33.5-35.1; respiratory siphon 30.6-30.9; width of head 3.42-3.42; width of an eye 1.14-1.15; Etymology interocular space 1.14-1.12; anterior width of prono- Recta (Latin, adjective, feminine form of rectus, tum 2.60-2.63; humeral width of pronotum 4.28- meaning straight) refers to the paramere having a 4.25; length of pronotum 8.9-8.8; length of anterior more parallel body than its closest relative R. incisa. lobe of pronotum 5.7-5.8; length of posterior lobe 3.2-3.0. Fore leg: mean value of length of distal part Ranatra sterea sp. n. of femur 5.0, proximal part of femur 6.6. Dimensions (figs. 3, 6, 11, 21, 22, 26, 47) of leg segments see table 1. Colour. – More or less uniformly medium brown, Type material. – Holotype (NMSC), female (half of except for anterior lobe of pronotum somewhat lighter siphon and left hind femur broken off), CHINA: Yun- than posterior lobe. Legs not annulated except for dis- nan, Baoshan (25˚1’N, 99˚1’E) Irawaddy basin, tal dark brown to blackish rings on segments of mid- Gaoligong mountain, 28.xi.2000, leg. H. H. Tan dle and hind legs. Structural characteristics. – A com- (THH0108). – Paratype, same data as holotype 1 paratively stout species (fig. 3). Lorum higher than (NCTN). clypeus, clypeoloral sulcus distinct and deep. Vertex Type locality. – Xishuangbanna in Yunnan low, not raised above eyes, without tubercle between Province, China. eyes (fig. 6). Eye width equal to interocular space. An- tenna as in fig. 11. Anterior margin of pronotum not

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raised, anterior lobe of pronotum slightly less than margin not incised and the female operculum always twice as long as posterior lobe, transverse groove dis- distinctly reaching beyond the apex of abdomen. Fi- tinct, sublateral swellings caudally on pronotum dis- nally R. linearis, which is of the same size and at first tinct but not prominent. Scutellum 1.3-1.5 times as sight looks similar, has the prosternum usually red- long as broad, dorsal surface smooth with a pair of dish, the metasternum not emarginate. Females of R. shallow depressions in caudal half. Prosternum linearis have the operculum narrower dorsoventrally, smooth, anterior lobe in posterior half with a tomen- with a smooth dorsal margin and an acute, slightly re- tose ridge of short clubby bristles, anterior half with a curved tip. few of such bristles in a row; prothoracic pit wide and shallow. Mesosternum smooth. Metasternum with a Distribution pair of sublateral grooves and a blunt median keel in Only known from type locality – Yunnan, South- posterior half; posterior margin distinctly emarginate west China. (fig. 26). Smallest distance between middle coxae 1.2 times the smallest distance between hind coxae. Mem- Etymology brane reaching just beyond posterior margin of sixth Sterea (Greek, adjective, feminine form of stereos, tergite (fig. 47). Sternite VI dorsoventrally expanded meaning robust) refers to the solid build of this caudally. Ventral margin of laterotergite VII and basal species. third of respiratory siphon with dense pilosity, anteri- orly on siphon long becoming shorter and less promi- Doubtful records nent caudally. Fore femur distinctly longer than pronotum, with a single, well developed tooth, ap- The following two species, in our opinion, do not proximately in apical two fifth, and a subapical inci- occur in China. As we have no opportunity to check sion without a small additional tooth (fig. 21-22). the specimens identified by Hoffmann, we include Width of fore femur measured from dorsal margin to them in this paper to facilitate further study. apex of tooth subequal to widest part of femur proxi- mally. Hind femur just reaching the base of sternite VI Ranatra filiformis Fabricius in paratype to basal fifth of sternite VI in holotype. (figs. 13, 19, 20, 37) Operculum in ventral view elongate with a promi- nent median keel with a fringe of sparse long bristles Ranatra filiformis Fabricius, 1790: 228. along the dorsal margin of the keel; in lateral view Ranatra filiformis; Lansbury 1972a: 321 (redescription, syn- boat-shaped with a comparatively blunt apex, just onymy); Polhemus, 1995: 18 (distribution). reaching the base of respiratory siphon (fig. 47); dor- sal margin of operculum serrate. Material examined. – INDIA: Karikal, iii.1960, leg. P.S. Male. – Unknown. Nathan 4 (RMNH); Karikal, xi.1961, leg. P.S. Nathan, 1 (RMNH); Pondicherry, Tranquebar, v.1952, leg. P.S. Nathan, 1 (NCTN); Pondicherry, Karikal, Kurukbagarum, iii.1962, Comparative notes leg. P.S. Nathan, 3 1 (NCTN). This species does not fit into any of the species groups defined by Lansbury (1972a). R. sterea shares Diagnosis the emarginate caudal margin of the metasternum Body length 23-28 mm, length of respiratory and the posteriorly dorsoventrally dilated sternite VI siphon 21-23 mm. Colour more or less uniformly pale with the species of the R. gracilis-group of Lansbury brown, with anterior lobe of prothorax a shade darker (1972a). Species of this group have the vertex raised than the rest of body. Legs not annulated except for above the eyes with a distinct tubercle, the respiratory distal dark brown to blackish rings on segments of siphon usually short, about one third of body length middle and hind legs. Vertex dorsally between eyes and the female operculum usually extending beyond raised, sometimes into a dorsal point suggesting a ves- the apex of abdomen. The exception is R. sulawesii tigial tubercle; eye width slightly less than interocular Nieser & Chen, 1991, which is about the same size as space. Antenna as in fig. 13. Transverse groove on pro- R. sterea, has a long siphon and the female operculum thorax shallow but well visible; anterior lobe of prono- not extending beyond the apex of abdomen. Apart tum smooth or with a pair of poorly developed shal- from having a tuberculate vertex, R. sulawesii differs low longitudinal impressions which are converging from R. sterea by having a longer and more slender posteriorly; length of pronotum 0.95-1.10 times the pronotum with a deep pronotal pit, and the respira- length of fore coxa. Length of fore coxa two thirds or tory siphon longer than the body. The other group of slightly less the length of fore femur; fore femur with species with a single prominent tooth centrally on two large teeth which are two fifth of fore femoral fore femur, the R. elongata-group of Lansbury length removed from distal apex of femur (figs. 19- (1972a), have a raised metasternum with its caudal 20). Prosternum with a median longitudinal keel ex-

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43

44

45

46

47

Figs. 43-47. Ranatrinae females, apex of abdomen in lateral view (scale 1 mm). – 43, Cercotmetus brevipes; 44, C. asiaticus; 45, Ranatra lansburyi, paratype; 46, R. incisa, paratype; 47, R. sterea, paratype.

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tending to level of transverse groove. Metasternum terior lobe, not extending onto posterior lobe; length not elevated, with a pair of shallow submarginal longi- of pronotum 0.9-1.0 times the length of fore coxa. tudinal impressions, not emarginate caudally. Length of fore coxa about two thirds the length of fore Male. – Paramere as in fig. 37. femur; fore femur with two large teeth which are two Female. – Genital operculum not extending be- fifth of fore femoral length removed from distal apex yond the apex of abdomen. of femur. Prosternum with a median longitudinal keel. Metasternum somewhat elevated, not emar- Discussion ginate caudally, with low blunt lateral ridges and a The occurrence of this species in China has to be blunt low median ridge in posterior half. confirmed. Lansbury (1972a) doubts its occurrence Male. – Paramere as in fig. 38. outside the Indian subcontinent and Sri Lanka. Polhe- Female. – Genital operculum not extending be- mus (1995) states that the habitus figure given by yond apex of abdomen. Hoffmann (1930b), based on specimens from Guang- dong, refers to R. filiformis. We have taken measure- Distribution ments from the figure in Hoffmann (1930b), they Recorded from most areas of China with the ex- match equally good with measurements of R. filiformis ception of the northeast and northwest (Hoffmann, as with those of R. recta sp. n. Hoffmann (1930b) 1941; Polhemus, 1995). Indonesia and Malaysia. states that his R. filiformis is very common in Guang- In view of the results of Lansbury (1972a), the dis- dong which would indicate a species of the R. longipes- covery that Chinese specimens from Taiwan and complex. Therefore we agree for the moment with Guangdong belong to R. incisa sp. n. and R. recta sp. Lansbury (1972a) that R. filiformis does not occur in n, and study of specimens of R. ‘longipes’ from various China, and assume that the R. filiformis of Hoffmann regions by the first two authors, we assume that the (1930b, 1941) actually belongs to R. recta. true R. longipes is restricted to Malaysia and the Sun- daland part of Indonesia. Consequently the Chinese Distribution records will belong to different species. See also the Nepal, India, Sri Lanka (Lansbury 1972a). comparative notes under R. recta.

Ranatra longipes Stål ACKNOWLEDGEMENTS (figs. 2, 38) Our sincere thanks are due to W. Hogenes (ZMAN), Ranatra longipes Stål, 1861: 203. J. van Tol (RMNH), and C. M. Yang (NMSC) for loan- Ranatra longipes; Hoffmann 1941: 57 (distribution in Chi- ing material under their care; to Chuan-ren Li (Hubei, na). Ranatra longipes longipes; Lansbury 1972a: 332 (redescrip- China), J.-R. Wu (Taiwan, China) for collecting the tion); Polhemus 1995: 18 (distribution). aquatic bugs and making them available for study. We are also indebt to G. Keijl (RMNH) for helping with the Material examined. – INDONESIA: Java: Java, v.d. Hoeven, literature. We are particularly grateful to J. van Tol, E. 1 (RMNH); Buitenzorg (= Bogor), v.d. Hoeven, 1 J. van Nieukerken, J. Huijbregts (RMNH) and H. (RMNH); Batavia (= Jakarta), 1882, Semmelinck, 1 Zettel (Natural History Museum in Vienna, Austria) (RMNH); Garut, 1928, W.C. v. Heurn, 1 (RMNH); Sukabu- for critical reading. and various suggestions to improve mi, ix.1924, le Moult, 2 (RMNH); Wangen (= Blawi) River, from Lamongan downstream, lowland stream, partly dried the manuscript. up with wide margins of soft mud, nearly stagnant, turbid due to suspended mud and algae, much Eichornia, REFERENCES 2.vii.1994, leg. N. Nieser, 1 3 teneral (NCTN). Bali: Road Sanur-Tohpati, km 4, small pond, water turbid light brown, Amyot, C. J. B. & A. Serville, 1843. Histoire naturelle des bottom clay/loam, some overhanging marginal vegetation, insectes. Hémiptères. – Librairie Encyclopédique de depth of sample point up to 0.2 m, 11.vii.1983, leg. N. Roret, Paris: lxxvi + 675 pp. Nieser, 10 4 (NCTN). Aukema, B. & C. Rieger (eds.), 1995. Catalogue of the Het- eroptera of the Palaearctic Region, 1. Enicocephalomor- pha, Dipsocoromorpha, Nepomorpha, Gerromorpha Diagnosis and Leptopodomorpha – The Netherlands Entomologi- Body length 20-27 mm, length of respiratory cal Society, Amsterdam, xxvi + 222 pp. siphon 14-25 mm (fig. 2). Colour uniformly pale Chen, P.-p., N. Nieser & H. Zettel, in press. The aquatic and semiaquatic bugs of Malesia and adjacent areas (In- brown, legs not annulated. Vertex distinctly raised secta- Heteroptera: Nepomorpha & Gerromorpha). – above eyes, in some specimens with a vestigial tubercle Fauna Malesiana Handbook. between eyes; eye width subequal to interocular space. Cloarec, A., 1972. Activité respiratoire de Ranatra linearis Transverse groove on prothorax well developed; (Insecte, Hétéroptère aquatique). – Bulletin de la Société pronotum with a poorly developed median keel on an- Zoologique de France 97: 115-124.

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True bugs of the world Keffer, S. L. & J. T. Polhemus, 1999. A new species of Cer- (Hemiptera: Heteroptera). – Cornell University Press, lo- cotmetus Amyot & Serville (Heteroptera: Nepidae) from cation, xii + 336 pp. Laos. – Journal of the New York Entomological Society Scott, J., 1874. On a collection of Hemiptera-Heteroptera 107: 28-30. from Japan. Descriptions of various new genera and Lansbury, I., 1972a. A review of the Oriental species of Rana- species. – Annals and Magazine of Natural History (4) tra Fabricius (Hemiptera-Heteroptera: Nepidae). – Trans- 14: 426-452. actions of the Royal Society of London 124: 287-341. Stål, C., 1861. Nova methodus familias quasdam Lansbury, I., 1972b. A revision of the genus Telmatotrephes Hemipterorum disponendi. – Öfversigt af Kungliga Stål (Hemiptera-Heteroptera, Nepidae). – Zoologica Vetenskapsakademiens Förhandlingar 18: 195-212. Scripta 1: 271-286. Stål, C., 1866. Hemiptera Africana 3 [1865]. – Nordste- Lansbury, I., 1973. 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Systema Naturae. Editio decima, refor- Received: 27 January 2004 mata. – Salvii, Holmiae, v + 824 pp. Accepted: 7 April 2004

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