Dtsch. Entomol. Z. 60 (2) 2013, 221–230 / DOI 10.1002/mmnd.201300028
The Berothidae of Taiwan (Neuroptera: Neuropterida)
Ulrike Asp ck*,1,2, Xingyue Liu3 and Horst Asp ck4
1 Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, 1010 Vienna, Austria. 2 Department of Integrative Zoology, University of Vienna, 1090 Vienna, Austria. 3 Department of Entomology, China Agricultural University, Beijing 100193, China. 4 Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University of Vienna, Kinderspitalgasse 15, 1095 Vienna, Austria.
Abstract
Received 17 June 2013 Only two species of the family Berothidae have been recorded from Taiwan: Isoscelip- Accepted 24 June 2013 teron puncticolle Nav s, 1911, and Isoscelipteron formosense (Kr ger, 1922). Based on Published 25 November 2013 our current findings, two new species are described and illustrated. The Berothidae are thus represented in Taiwan by two genera, Isoscelipteron Costa, 1863, with the two aforementioned species and Berotha Walker, 1860, with two new species: Berotha spe- Key Words tana sp. n. and Berotha taiwanica sp. n. Furthermore, the hitherto unknown female of I. formosense is described and illustrated for the first time. The relationships to other Berotha species of Berotha and Isoscelipteron respectively are discussed. A key to the genera Isoscelipteron and species of Berothidae of Taiwan is provided. The records of Berothidae of Taiwan new species are shown in a map. Probably all four species are endemic to the island.
Introduction Materials and methods
The Berothidae, a small neuropteran family comprising The specimens examined in the present study comprise pinned mate- about 120 known species representing six subfamilies, rial convened from own collecting activities and borrowed material are scattered around the globe (U. Asp ck & Nemesch- from diverse collections: Natural History Museum Vienna (NHM), Senckenberg, Deutsches Entomologisches Institut M ncheberg kal 1998, U. Asp ck, Randolf & Zimmermann 2013). (SDEI), Museum f r Naturkunde der Humboldt-Universit t zu Berlin In Taiwan so far only two species of the family have (ZMB), coll. Bo Tjeder, now Museum of Zoology and Entomology, been recorded, both belong to the genus Isoscelipteron Lund University (MLU), Zoologisk Museum Kopenhagen (ZMK), Costa, 1863: Isoscelipteron puncticolle Nav s, 1911, Mus um National D’Histoire Naturelle, Paris (MHN); see acknowl- and Isoscelipteron formosense (Kr ger, 1922). A study edgements. Preparations of the genital segments were made in the of a number of specimens of Berothidae from Taiwan, usual manner with KOH. The cleared genitalia were preserved in gly- housed in various collections, revealed two new species, cerin. The terminology of the genitalia follows that of H. Asp ck et al. (1980) and U. Asp ck & H. Asp ck (2008). both belonging to the genus Berotha Walker, 1860. The present paper gives an overview of our current knowl- edge of the Berothidae of Taiwan, as well as descrip- tions of the new species.
Genera and species of Berothidae found in Taiwan
Key to the Berothidae of Taiwan
1. Forewingalongthepterostigmadistinctlyconvex(Fig.8)...... ( Berotha) 2 – Forewingalongthepterostigmainconspicuous(Fig.15)...... ( Isoscelipteron) 3 2. Sternite 8 of male with median processus, tergite 9 þ ectoproctofthefemaleventrallybroad,nottapering(Figs4,5)...... B. spetana
* Corresponding author, e-mail: [email protected]
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– Sternite 8 of male without processus, tergite 9 þ ectoproctofthefemaleventrallytapering(Fig.10)...... B. taiwanica 3. Gradatesofhindwingendingbelowpterostigma(Fig.15)...... I. formosense – Gradatesofhindwingendingproximallyofpterostigma...... I. puncticolle
Genus Berotha Walker, 1860 our books. The specific epithet spetana is an adjective in the nominative feminine singular. Typus generis. Berotha insolita Walker, 1860. Traditionally, the genus Berotha has been a mixtum Locus typicus. Taiwan, Huisun. compositum of various species of Berothidae having Material studied. Holotype, male “Berotha spetana U. Asp ck et X. different degrees of relationships (Nav s 1929). The Liu et H. Asp ck Holotypus male / Taiwan, Prov. Nantou, Huisun, taxonomic clarification and characterisation of Berotha, 24 0503900 N 121 0200200 E, 800 m, 11 Mai 1999, T99/1, H. & U. As- as well as recognition of the genus as an Oriental ele- p ck leg.”, coll. NHM. ment, were published almost half a century later (U. Paratypes. 1 male “Kosempo Formosa [Taiwan: Kaohsiung County, Asp ck 1983). Chiasien (= Kosempo), 23 040 N, 120 340 E] H. Sauter, 1911/7.VII. / Petersen det. / Berotha puncticollis Nav. det. Esb.Peters. / formosensis Presently nine species are assigned to the genus. Kr g.”, coll. SDEI; 1 female “(Tainan) Formosa [23 0804600 N B. insolita Walker, 1860 (India); B. indica (Brauer, 120 1500500 E] / Frawalkeria puncticollis Nav. P. Nav s S.J. det. / syn- 1865) (Sri Lanka); B. piepersii Van der Weele, 1904 type”, coll. SDEI; 1female “Rengeti Formosa [Taiwan: Nantou 0 0 (Singapore and Indonesia: Java); B. borneensis Nav s, County, Lienhuachi (= Rengeti), 23 53 N, 120 53 E] 29.III.27 / Coll. Esben Petersen”, ZMK. 1912 (Borneo: Sarawak); B. insulana U. Asp ck & H. Asp ck, 1981 (Indonesia: Riau Archipelago); B. banna- na (Yang, 1986) (China: Yunnan); B. zhejiangana Yang Description & Liu, 1995 (China: Zhejiang); B. baminana Yang & Liu, 1999 (China: Fujian); and B. chouioi Yang & Liu, Length of fore wing 11–12 mm (male), 12–12.5 mm 2002 (China: Hainan) (U. Asp ck & H. Asp ck 1981, (female). Fore wings prominently falcate. Fore wing as U. Asp ck 1983, Yang 1986, Yang et al. 1995, 1999, well as hind wing without scale like hairs. 2002, Oswald 1998). Scapus approximately as long as the following 7 to 8 flagellomeres. Antennae yellowish with yellow hairs, scapus with reddish brownish dots. Head and pronotum Berotha spetana sp. n. yellow with reddish-brown dots, pronotum with a thin Figures 1–6, 20 brown median line. Hairs of pronotum yellowish dorsally, brownish ventrally. Fore wing distinctly convex in the Derivatio nominis: The new species is cordially dedi- area of the pterostigma. Wing membrane hyaline yellow- cated to Dr. Franz Speta, Univ. Dozent [associate pro- ish, pterostigma reddish-brownish dotted, with hyaline fessor] of Systematic Botany at the University of Salz- parts in the middle. Longitudinal veins yellowish, brown- burg and former Director of the Biologiezentrum Linz ish dotted, crossveins brownish. Hind wing with incon- on the occasion of his 70th birthday (22 December spicuous pterostigma, longitudinal veins yellowish. 2011) as a token of our appreciation for his outstanding scientific work and our gratitude for his decisive invol- Male genitalia (Figs 3–4). Sternites 7 and 8 with vement and expertise in the publication of several of ridges, sternite 8 caudally with median digitiform pro-
Figure 1. Berotha spetana sp. n., male, holotype. Taiwan, Huisun. Length of fore wing 12 mm.
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Figure 2. Berotha spetana sp. n., female, paratype. Taiwan, Tainan. Length of fore wing 12 mm. – A: Analis; b: basal piece of Ma; Cua: Cubitus anterior; Cup: Cubitus posterior; Ma: Media anterior; Mp: Media posterior; Pt: pterostigma; R: Radius; Rs: Radius sector; Sc: Subcosta.
Figures 3–4. Berotha spetana sp. n., male, holotype. Taiwan, Huisun. Genital segments, lateral (3) and ven- tral (4). – e: ectoproct; g: gonarcus (fused gonocoxites 11); gx9: gonocoxite 9; hi: hypandrium internum; p: fused parameres (complex of fused gonocoxites, gona- pophyses, gonostyli 10); S 7–9: sternites 7, 8, and 9; T7–9: tergites 7, 8, and 9.
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Figures 5–6. Berotha spetana sp. n., female, paratype. Taiwan, Tainan. Genital segments, lateral (5), and ventral (6). – b: bursa copulatrix; e: ectoproct; hc: hy- pocauda; gp7: gonapophyses 7; gx9: gonocoxites 9; pd: pudiculum (gonapo- physes 8); sg: subgenitale (fused gonocoxites 8); r: spermatheca; S 7: sternite 7; T7–9: tergites 7, 8, and 9.
cessus. Tergite 8 surrounding spiraculum, tergite 9 þ coiled ductus, small but strongly sclerotised globular ectoproct shell like. Trichobothria very indistinct. Ster- element. nite 9 of paired appearance, comprising lateral triangu- Differentiation. Due to the male genitalia, B. spetana lar sclerites, equipped with dorsoventral and lateral sp. n. is most closely related to B. insulana U. Asp ck & ridges, both sclerites connected ventrally by a sclero- H. Asp ck, 1981, from the Riau Archipelago (Indone- tised bar, apices with two and three little teeth like pro- sia), which can be differentiated from B. spetana by the cessus respectively. Gonocoxites 9 with hooked apex. much shorter bristle-bow of the gonocoxite-complex 10, Gonocoxites 11 (gonarcus) forming a small sclerotised the broad incision of sternite 9 in the male, and by the bow, fused with gonocoxites 9, whereby the complex ventrally tapering tergite 9 þ ectoproct in the female. comprises paired rods which reach into segment 8. The differentiation from B. taiwanica sp. n., see there. Complex of fused gonocoxites, gonapophyses and go- nostyli 10 (parameres) with a large basal sclerite and a Distribution (Fig. 20). B. spetana sp. n. is known from long bow formed by long bundled bristles, reaching the type locality and, in addition, from the apparently into segment 7 cephally and ending as a bundle of free traditional collecting spot Chiasien (Kosempo), there bristles terminally. Hypandrium internum large. syntoptic with I. formosense. B. spetana sp. n. is most Female genitalia (Figs 5–6). Sternite 7 mostly re- probably endemic to Taiwan. duced, gonocoxites 7 forming caudally domed sclerites, gonapophyses 7 forming huge smooth sclerite discs. Subgenitale (gonocoxite 8) ribbon-like, laterally with Berotha taiwanica sp. n. a sclerotised ridge, ventrally with paired processus, of Figures 7–12, 20 semicircular shape. Gonapophyses 8 forming a tiny unpaired sclerite (pudiculum). Tergite 9 þ ectoproct Derivatio nominis: The specific epithet is an adjective ventrally broad, not tapering. Bursa copulatrix-sper- in the nominative feminine singular; it refers to the dis- matheca complex with broad atrium bursae, loosely tribution which is probably restricted to Taiwan.
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Figure 7. Berotha taiwanica sp. n., fe- male, holotype. Taiwan, Dalin (= Taihor- in). Length of fore wing 9.2 mm.
Figure 8. Berotha taiwanica sp. n., male, paratype. Taiwan, Dalin (= Taihorin). Length of fore wing 9 mm.
Locus typicus. Taiwan, Dalin (= Taihorin). H. Sauter s. g. / Berotha indica Brau. P. Nav s S. J. Material studied. Holotype, female “Formosa Taihorin [Taiwan: Chiayi det. / MUSÞUM PARIS”, coll. MNP; 1 female “For- County, Dalin (= Taihorin), 23 350 N, 120 280 E] 11.1911 Saut. / photo mosa Taihorin [Taiwan: Chiayi County, Dalin (= Tai- compared with type of B. insolita Walk. / Berotha indica Brauer m / Coll. horin), 23 350 N, 120 280 E] V.10. H. Sauter S. G. / Esben-Petersen / Berotha insolita Walk. det. Esben.Petersen”, coll. ZMK. Berotha indica Br. / Berotha indica Brau. P. Nav s S. J. Remark. The only available male specimen has da- det. / Zool Mus. Berlin”, coll. ZMB; 1 female (genital maged genital segments, therefore we selected a female sclerites embedded in Canada balsam) “Sokutsu Formo- specimen as the holotype. sa [Taiwan: Kaohsiung County, near Chiasien, Hsiaolin (= Sokutsu), 23 0604200 N, 120 3605900 E] H. Sauter Paratypes. 1 male “Formosa Taihorin [Taiwan: Chiayi V.1912 / Berotha indica m det. Esben-Petersen / 19”, County, Dalin (= Taihorin), 23 350 N, 120 280 E] IV.10. coll. SDEI.
Figure 9. Berotha taiwanica sp. n., fe- male, paratype. Taiwan, Dalin (= Taihor- in). Length of fore wing 9 mm.
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Description
Length of fore wing 9 mm (male), 9–9.2 mm (female). Fore wings prominently falcate. Fore wing as well as hind wing without scale-like hairs. Scapus approximately as long as the following 6 to 7 flagellomeres. Antennae yellowish with yellow hairs, scapus with reddish-brownish dots. Head and pronotum yellow with reddish-brown dots, pronotum without dots medially. Hairs of pronotum yellowish dorsally, brown- ish ventrally. Fore wing distinctly convex in the area of the pterostigma. Wing membrane hyaline yellowish, pterostigma reddish brownish, with hyaline window in the middle. Longitudinal veins yellowish, brownish dotted, crossveins brownish. Hind wing with inconspic- uous pterostigma, with reddish-brown veins; longitudi- Figure 12. Berotha taiwanica sp. n., male, paratype, Taiwan, nal veins yellowish. Dalin (= Taihorin). Female genitalia (Figs 10–11). Sternite 7 mostly re- duced, gonocoxites 7 forming caudally domed sclerites, Male genitalia (Fig. 12). Severely damaged. Appar- gonapophyses 7 as huge smooth sclerite discs (visible ently very similar to B. insolita from India (see U. As- particularly in the drawing of the ventral aspect). Sub- p ck 1983, Figs 4–7). Tergite 8 surrounding spiracu- genitale (= fused gonocoxites 8) ribbon-like laterally, lum, sternite 8 inconspicuous. Tergite 9 þ ectoproct ventrally with paired inconspicuous hooks. Gonapo- ventrolaterally with flat incision between both; tricho- physes 8 forming a tiny sclerite (pudiculum). Tergite 9 bothria indistinct. Sternite 9 small, inconspicuous. Go- ectoproct ventrally tapering. Bursa copulatrix-sper- þ nocoxites 9 with small hooked apex. Gonocoxites 11 matheca complex with loosely coiled ductus, small but (gonarcus) forming a small sclerotised bow. Complex strongly sclerotised globular element (This globular ele- of fused gonocoxites, gonapophyses and gonostyli 10 ment shows an impression, which is certainly artificial (parameres) mostly damaged, but apparently extremely and has therefore not been indicated in the drawing.) tiny, the bundled bristles almost invisible. Hypandrium internum large. Differentiation. Due to female and male genitalia, B. taiwanica sp. n. is most closely related to B. insolita Walker, 1860, from India and B. indica (Brauer, 1865), from Sri Lanka, from both it is differentiated by the much shorter bristle-bow of the gonocoxite-gonapo- physes-gonostylus-complex 10 (parameres) in the male, by the lack of traces of sternite 7, and lack of sclero- tised ridges on the subgenitale, and a ventrally less ta- pering tergite 9 þ ectoproct in the female. B. spetana n.sp. is differentiated prima vista by the median proces- sus of sternite 8 in the male and by the ventrally broad and not tapering tergite 9 þ ectoproct in the female. Distribution (Fig. 20). B. taiwanica sp. n. is known from the type locality and from Hsiaolin (= Sokutsu), and is most probably endemic to Taiwan.
Genus Isoscelipteron Costa, 1863 Typus generis. Isoscelipteron fulvum Costa, 1863. The disorder concerning the coverage of the genus, to which Nav s (1929) contributed essentially, could be clarified in several steps by U. Asp ck (1983) and U. Asp ck & H. Asp ck (1980, 1981 and 1991; a detailed synonymy list is presented in H. Asp ck et al. 2001). At present, the genus comprises 14 species: Figures 10–11. Berotha taiwanica sp. n., female, holotype, Taiwan, Dalin (= Taihorin), genital segments, lateral (10) and I. fulvum Costa, 1863 (East Mediterranean and Middle ventral (11). East); I. pectinatum (Nav s, 1905) (China); I. puncti-
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Figure 13. Isoscelipteron formosense (Kr ger, 1922), male. Taiwan, Chiasien (= Kosempo). Length of fore wing 12.5 mm. colle Nav s, 1911 (China: Taiwan); I. rufum (Nav s, Isoscelipteron formosense (Kr ger, 1922) 1912) (Australia: Queensland); I. nicobaricum (Nav s, Figures 13–19, 20 1912) (Nicobar Islands); I. okamotonis (Nakahara, 1914) (Japan); I. formosense (Kr ger, 1922) (China: Acroberotha formosensis Kr ger, 1922 (original description): Nav s Taiwan); I. tonkinense (Kr ger, 1922) (Vietnam); I. le- 1929 (description); Rousset 1968 (description, figs of wings and veri (Kimmins, 1951) (Solomon Islands); I. glaserellum (uncleared) terminalia, taxonomy, lectotype designation). Isoscelipteron formosense (Kr ger, 1922): U. Asp ck & H. Asp ck H. Asp ck, U. Asp ck & H lzel, 1979 (Western Medi- 1980 (redescription, figures of wings and of male genital sclerites terranean); I. philippinicola U. Asp ck & H. Asp ck, of lectotype); U. Asp ck & H. Asp ck 1981 (listed); Yang et al. 1991 (Philippines); I. borneole U. Asp ck & H. As- 1995 (taxonomic comment). p ck, 1991 (Borneo); I. dictyophilum Yang & Liu, 1995 (China: Zhejiang); I. eucallum Yang & Liu, 1999 (Chi- Locus typicus. Taiwan, Chiasien (= Kosempo). na: Fujian) (U. Asp ck 1983, U. Asp ck & H. Asp ck Material studied. 1 male “Kosempo Formosa [Taiwan: Kaohsiung 0 0 1980, 1981, 1991, Yang et al. 1995, 1999, H. Asp ck County, Chiasien (= Kosempo), 23 04 N, 120 34 E] H. Sauter, 1911 / 7.XII / Petersen det. / formosensis Kr g.”, coll. SDIE; 1male “Formo- et al. 2001). sa (Kosempo) [Taiwan: Kaohsiung County, Chiasien (= Kosempo), 23 040 N, 120 340 E] Sauter. / Berotha puncticollis Nav. / Zool. Mus. Berlin”, coll. ZMB; 1 male “Kosempo Formosa [Taiwan: Kaohsiung Isoscelipteron puncticolle Nav s, 1911 County, Chiasien (= Kosempo), 23 040 N, 120 340 E] H. Sauter, 1911 / Isoscelipteron puncticolle Nav s, 1911 (original description): U. As- Petersen det. / 7. XII”, coll. SDEI; 2 males “Kosempo Formosa [Tai- 0 0 p ck & H. Asp ck 1980 (comment). wan: Kaohsiung County, Chiasien (= Kosempo), 23 04 N, 120 34 E] Berotha puncticollis (Nav s, 1911): Nav s 1912 (nomenclature); Kr - H. Sauter, 1911 / Petersen det. / 7. XII / Berotha puncticollis Nav. ger 1922 (monograph); Stange & Wang 1998 (taxonomic com- det. Esb. Peters.”, coll. SDEI; 3 males “Kosempo Formosa [Taiwan: 0 0 ments, photograph, records). Kaohsiung County, Chiasien (= Kosempo), 23 04 N, 120 34 E] H. Frawalkeria puncticollis (Nav s, 1911): Nav s 1929 (monograph). Sauter, 1911 / Petersen det. / 7. XII.”, coll. SDEI; 1 male “Kosempo Formosa [Taiwan: Kaohsiung County, Chiasien (= Kosempo), Locus typicus. Taiwan, Tainan. 23 040 N, 120 340 E] H. Sauter, 1911 / 7. XII. Berotha puncticollis So far it has not been possible to examine the type Nav. det. Esb. Peters. / Acroberotha formosensis Kr g. P. Nav s S. J. of this species; we even do not know if it still exists. It det.”, coll. SDEI; 1 male “Kosempo Formosa [Taiwan: Kaohsiung 0 0 cannot be excluded that I. puncticolle and I. formosense County, Chiasien (= Kosempo), 23 04 N, 120 34 E] H. Sauter, 1911 / are the same species. In this case I. formosense would Petersen det. / Berotha puncticollis Nav. det. Esb. Peters. / 22. XII”, coll. SDEI; 1 male “Kosempo Formosa [Taiwan: Kaohsiung County, become a younger synonym. The photograph published Chiasien (= Kosempo), 23 040 N, 120 340 E] H. Sauter 1911 / 7. XII / by Stange & Wang (1998) shows apparently a specimen Petersen det. / Berotha puncticollis Nav. det Esb. Peters.” coll. Bo Tje- of that species which we know to be I. formosense (see der, now MLU; 1female “Kosempo Formosa [Taiwan: Kaohsiung below). County, Chiasien (= Kosempo), 23 040 N, 120 340 E] H. Sauter, 1911 /
Figure 14. Isoscelipteron formosense (Kr ger, 1922), male. Taiwan, Chiasien (= Kosempo). Length of fore wing 13 mm.
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Figure 15. Isoscelipteron formosense (Kr ger, 1922), male. Taiwan, Chiasien (= Kosempo). Length of fore wing 12.2 mm.
Figure 16. Isoscelipteron formosense (Kr ger, 1922), female. Taiwan, Chia- sien (= Kosempo). Length of fore wing 10.1 mm.
7. XII. Berotha puncticollis Nav. det. Esb. Peters. / label with male Nantou Hueisun Exp. Forest [24 0503900 N, 121 0200200 E] 24.–29. X. symbol / Acroberotha formosensis Kr g. P. Nav s S. J. det. female 1998, 650 m leg. Mey & Speidel”, coll. ZMB. symbol”, coll. SDEI; 1male “Kosempo Formosa [Taiwan: Kaohsiung County, Chiasien (= Kosempo), 23 040 N, 120 340 E] / Coll. Esben- Description of the hitherto unknown female Petersen / Acroberotha formosensis Krg. Underside: Berotha puncti- The only female available from the type locality collis Nav”, coll. ZMK; 1 male “Sokutsu Formosa [Taiwan: Kaoh- (Fig. 16) is similar in shape and appearance to the siung County, near Chiasien, Hsiaolin (= Sokutsu), 23 0604200 N, 120 3605900 E] H. Sauter V. 1912 / Petersen det. / formosensis Kr g.”, males (Figs 13–15). There are, however, single scales, coll. SDEI; 1 male “Petersen det. / Sokutsu Formosa [Taiwan: Kaoh- scattered mainly along longitudinal veins in the proxi- siung County, near Chiasien, Hsiaolin (= Sokutsu), 23 0604200 N, mal third of the hind wing, but it seems that most of 120 3605900 E] H. Sauter V. 1912”, coll. SDEI; 1 female “Taiwan, the scales are lost. Length of fore wing 10.1 mm.
Figure 17. Isoscelipteron formosense (Kr ger, 1922), female, Taiwan, Huei- sun. Length of fore wing 12.5 mm.
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set of scale like hairs in the basal third of the hind wing (Fig. 17). Unfortunately, for unknown reasons, the geni- tal segments of this species withstood the usual macera- tion, nevertheless we can compare the corresponding structures adequately. The bursa-copulatrix-spermatheca complex and the latero-ventral constriction of tergite 9 agree sufficiently to assign this specimen to I. formo- sense.
Discussion
The two genera, Isoscelipteron and Berotha recorded from Taiwan (with altogether four species) are widely distributed in East Asia and in other parts of the Pa- laearctic. Including the two new species from Taiwan, the genus Berotha now comprises 11 known species and is a fascinating example for a radiation occurring Figures 18–19. Isoscelipteron formosense (Kr ger, 1922), fe- male. Taiwan, Chiasien (= Kosempo), genital segments, lateral in the Oriental realm. (18) and subgenitale, ventral (19). Records of Berotha species are known from India, Sri Lanka, various parts of China, Singapore, and var- ious parts of Indonesia; the two species discovered in Female genitalia (Figs 18–19). Sternite 7 almost ob- Taiwan are most probably endemic to the island. B. spe- literated, gonocoxites 7 forming semicircular domed tana sp. n. is related to B. insulana from the Riau Ar- sclerites, gonapophyses 7 forming huge smooth sclerite chipelago, while B. taiwanica sp. n. is most closely re- discs. Subgenitale (fused gonocoxites 8) with sclero- lated to B. insolita and B. indica. This means that the tised ridge laterally, with paired lobes ventrally. Gona- two Berotha species so far found in Taiwan must be of pophyses 8 (pudiculum) inconspicuous. Tergite 9 þ ec- different geographic origin. From the morphological toproct latero-ventrally somewhat constricted. Bursa- characters of B. spetana n.sp., we tentatively conclude copulatrix-spermatheca complex comprising a basal that the species can be traced back to an old invasion, coil with broad lumen and a subsequent coil with thin- at least in the early Pleistocene. Due to the close rela- ner lumen, and with small but strongly sclerotised glob- tionship of B. taiwana to species occurring on the ular element. Both hypocaudae broken at base. Asian mainland, it may be assumed that it can be traced The only other available female is from Huisun back to a younger immigration. (Fig. 20), and is equipped with an apparently complete The genus Isoscelipteron presently comprises 14 spe- cies and has a distribution covering the Mediterranean, the Middle East, Nicobar Islands, East- and South East Asia (China, Taiwan, Japan, Vietnam), the Solomon Is- lands and the north of Australia (Queensland). Thus Isoscelipteron is distributed throughout three biogeogra- phical regions: the Palaearctic, the Oriental and the Australian region. Isoscelipteron formosense is closely related to I. tonkinense and can be regarded as having a young origin. During the Pleistocene the sea level dropped down repeatedly by 130 m (possibly even 140 m) compared to the present sea level. This led to an extension of the coast lines of the east of Asia by 600 up to 1000 km eastwards. Such events occurred at least one million years, 500,000 years, and 300,000 years B.P., and also during the last glacial period the sea level dropped by 120 m during the maximum of glaciation (ca. 18,000 years B.P.), which led to a broad connection of Taiwan with the mainland of China (Smith et al. 1995, Zhuo et al. 1998, Keally 2005, Liu et al. 2010). Nothing is known on the biology of the Berothidae Figure 20. Records of Berothidae in Taiwan. Circle: Berotha of Taiwan. As far as we know all specimens were taken spetana, quadrat: Berotha taiwanica, triangle: Isoscelipteron at light within arboreal biotopes in mountainous re- formosense, star: Isoscelipteron puncticolle. gions. No larvae of Berothidae have ever been found in
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Taiwan. It is hypothesized that Berothidae are generally 1997. Proceedings of the Sixth International Symposium on Neu- associated with termites. The complete life-cycle, ropterology, Helsinki, Finland, 13–16 July 1997. – Acta Zoologi- which comprises a non-feeding second instar with re- ca Fennica 209: 45–63. Asp ck, U., Randolf, S. & Zimmermann, D. 2013. Beaded lacewings duced legs and mouth parts, was successfully studied in – a new phylogenetic start-up (Insecta: Neuropterida: Berothidae) the American species Lomamyia latipennis Carpenter, [Talk]. In Kroh, A. et al. (eds). BioSyst.EU 2013, Global Sys- 1940 by Tauber & Tauber (1968). Whether the associa- tematics! 18–22 February 2013, Vienna, Austria. Abstract Vol- tion with termites and a hypermetamorphosis belong to ume: 16. the ground pattern of the family remains, however, to Keally, C. T. 2005. Japanese Pleistocene Landbridges and the Earliest be clarified. At any rate the biology of Berothidae is Watercraft. – Japanese Archaeology. [homepage on the internet] indeed a wide open field. Tama Cable Network, Tokyo, Japan. [Cited 21 July 2010.] Avail- able from URL: http://www.tnet.ne.jp/~keally/MiddlePalaeol/land- bridges.html Acknowledgements Kr ger, L. 1922. Berothidae. Beitr ge zu einer Monographie der Neu- ropteren-Familie der Berothiden. – Stettiner Entomologische Zei- We are much indebted to Prof. Yang Tsen-Tse (University of Tai- tung 83: 49–88. chung, Taiwan) who enabled us to carry out neuropterological field- Liu, X. Y., Asp ck, H., Hayashi, F. & Asp ck, U. 2010. New species work in Taiwan, grateful thanks also are due to his student Pan of the snakefly genus Mongoloraphidia (Raphidioptera: Raphidii- Chien-An who accompanied us in Taiwan and helped us in many dae) from Japan and Taiwan, with phylogenetic and biogeographi- ways. Specimens of Berothidae from Taiwan could be borrowed from cal remarks on the Raphidiidae of Eastern Asia. – Entomological several institutions: Cordial thanks go to Dr. Reinhard Gaedike, and Science 13: 408–416. Dr. Stephan M. Blank (Deutsches Entomologisches Institut Ebers- Nav s, L. 1911. N vropt res nouveaux de l’extr me Orient. – Russ- walde and M ncheberg, respectively), Dr. Roy Danielsson (Museum koe Entomologicheskoe Obozrenie [= Revue Russe d’Entomolo- of Zoology and Entomology Lund University), Dr. Kurt K. G nther gie] 11: 111–117. (Museum f r Naturkunde der Humboldt-Universit t zu Berlin), Prof. Nav s, L. 1912. Cris pidos y Hemer bidos (Ins. Neur.) nuevos cri- Niels P. Kristensen (Zoologisk Museum Kopenhagen), Dr. J. Legrand ticos. – Broteria 10: 98–113. (Mus um National D’Histoire Naturelle, Paris). A big thank you to Nav s, L. 1929. Monograf a de la familia de los Ber tidos (Insectos, Mag. Harald Bruckner (Natural History Museum, Vienna) for the ex- Neur pteros). – Memorias de la Real Academia de Ciencias Ex- cellent photographic documentation and for his competent graphical actas, Fisico-Quimicas y Naturales de Zaragoza 2: 1–107. assistance. Finally, we appreciate Dr. Dr. John Plant (Vienna) for lin- Oswald, J. 1998. Annotated catalogue of the Dilaridae (Insecta: Neu- guistic improvement of the manuscript. roptera) of the World. – Tijdschrift voor Entomologie 141: 115– 128. Rousset, A. 1968. Redescription de quelques types de B rothid s References (N vropt res Planipennes) d pos s au Mus um National d’His- toire Naturelle de Paris et remarques sur la syst matique de la Asp ck, H., Asp ck, U. & H lzel, H. (unter Mitarbeit von Rausch, famille. – Bulletin du Mus um National d’Histoire Naturelle, H.). 1980. Die Neuropteren Europas. Eine zusammenfassende Paris (2) 40: 275–291. Darstellung der Systematik, kologie und Chorologie der Neurop- Smith, A. G., Smith, D. G. & Funnell, B. M. 1995. Atlas of Mesozoic teroidea (Megaloptera, Raphidioptera, Planipennia) Europas. and Cenozoic Coastlines. Cambridge University Press, Cam- Goecke & Evers, Krefeld: 495 pp. and 355 pp. bridge: 99 pp. Asp ck, H., H lzel, H. & Asp ck, U. 2001. Kommentierter Katalog Stange, L. A. & Wang, H. Y. 1998. Guide book to insects in Taiwan. der Neuropterida (Insecta: Raphidioptera, Megaloptera, Neurop- 18. Neuroptera, Megaloptera, Raphidioptera. Hsu Hsin Books Ltd. tera) der Westpal arktis. – Denisia 2: 1–606. Co., Taipei, Taiwan: 278 pp. Asp ck, U. 1983. Das Genus Berotha Walker (Neuropteroidea: Plani- Tauber, C. A. & Tauber, M. J. 1968. Lomamyia latipennis (Neurop- pennia: Berothidae). – Annalen des Naturhistorischen Museums tera, Berothidae) life history and larval descriptions. – Canadian Wien 84/B: 463–478. Entomologist 100: 623–629. Asp ck, U. & Asp ck, H. 1980. Das Genus Isoscelipteron Costa, Yang, C. K. 1986. Thirty new species and four new genera of Neu- 1863 (Neuropteroidea: Planipennia: Berothidae). – Zeitschrift der roptera from Yunnan, and the family Nemopteridae new to China. Arbeitsgemeinschaft sterreichischer Entomologen 32: 65–74. – Acta Agriculturae Universitatis Pekinensis [= Peiching Nung Asp ck, U. & Asp ck, H. 1981. Weitere Untersuchungen an Berothi- Yeh Ta Hsueh Pao] 12: 153–166, 423–434. den: Berotha Walker, Isoscelipteron Costa und Asadeteva n.g. Yang, C. K. & Liu, Z. Q. 1999. Berothidae. In Huang, B. K. (ed.). (Neuropteroidea: Planipennia). – Zeitschrift der Arbeitsge- Fauna of insects in Fujian Province of China, Volume 3. Fujian meinschaft sterreichischer Entomologen 33: 1–14. Science Press, Fuzhou: pp. 100–102. Asp ck, U. & Asp ck, H. 1991. Zur Kenntnis des Genus Isoscelipter- Yang, C. K. & Liu, Z. Q. 2002. Neuroptera: Berothidae. In Huang, on Costa, 1863 (Neuropteroidea: Neuroptera: Berothidae: Berothi- F. S. (ed.). Forest insects of Hainan. Science Press, Beijing, China: nae). – Zeitschrift der Arbeitsgemeinschaft sterreichischer Ento- pp. 284–285. mologen 43: 65–76. Yang, C. K., Liu, Z. Q. & Yang, X. K. 1995. Neuroptera. In Wu, H. Asp ck, U. & Asp ck, H. 2008. Phylogenetic relevance of the genital (ed.). Insects of Baishanzu Mountain, eastern China. China For- sclerites of Neuropterida (Insecta: Holometabola). – Systematic estry Publishing House, Beijing: pp. 276–285. Entomology 33: 97–127. Zhuo, Z., Baoyin, Y. & Petit-Maire, N. 1998. Paleoenvironments in Asp ck, U. & Nemeschkal, H. L. 1998. A cladistic analysis of the China during the Last Glacial Maximum and the Holocene Opti- Berothidae (Neuroptera). In Panelius, S. (ed.). Neuropterology mum. – Episodes 21: 152–158.
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