Bull. Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A, 6: 23-31, March 31, 2008

New and additional specimens of giant from the Lower of (: : Palaeontinidae)

Kyoichiro UEDA

Kitakyushu Museum of Natural History and Human History, 2-4-1 Higashida, Yahatahigashi-ku, Kitakyushu 805-0071, Japan

（Received October 1, 2007; accepted January 20, 2008）

ABSTRACT ― Four new species of Palaeontinidae are described from the Lower Cretaceous of Araripe, Northeast Brazil: Parawonnacottella signifer n. sp., Colossocossus acutus n. sp., Baeocossus rotundatus n. sp., and Baeocossus muratai n. sp. Furthermore, a new complete adult specimen with wings is described for the species Parawonnacottella araripensis.

During the curation of the fossil collection of and closely situated medially; medial region of head pointed in Kitakyushu Museum of Natural History and Human History dorsal view; a deep furrow present on pronotum; mesothorax (KMNH), five specimens of Palaeontinidae from Araripe (Lower large, 15 mm long x 20 mm wide, at its widest point, bearing a Cretaceous) of Brazil were found and are described here. On the distinct medial ridge; abdomen 12 mm long with six discernible wing venation system I follow WhALLEY & JARzEmBOWSKI (1985) segments; posterior margin of each segment ornamented with and the names by KUKALOVá-PECK (1991) and DWORAKOWSKA minute punctures; ornamented region broadened towards terminal (1988) are added in the parentheses on each figures. All the segment. Fore wing missing antero-basal region between costa previously described species of Palaeontinidae from the Crato and R; cross vein CuA2-CuP distinct; traces of wing patterns Formation have recently been reviewed by MEnOn, HEADS & present alongside each vein, especially remarkable on apical

SzWEDO in MARTILL et al. (2007: 284-289). region beyond M2 and forming a narrow line curved inwardly. Hind wing narrow with distinct three veins; Rs branched from R

For all specimens the locality and horizon is Nova Olinda, basally from midwing; M1 evenly curved. Araripe Basin, N. E. Brazil, , Upper to Lower Albian (MAISEY, 1990). All scales in the figures are 10 Almost complete adult specimen preserved in dorsal aspect. mm. Dimension. Fore wing length 48 mm, width 21 mm; hind wing length 25 mm; wing expanse 95 mm; pronotum length 16 mm, SYSTEMATIC PALAEONTOLOGY width 21 mm; abdomen length 13 mm; total body length 30 mm. Superorder Depository. MURATA collection in KMNH. Auchenorrhyncha Suborder Cicadomorpha Remarks. This is the first record of P. araripensis in the almost Superfamily Cicadoidea complete condition. The small head and the closely situated eyes Family Palaeontinidae HAnDLIRSCh, 1906 are remarkable. The venation of hind wing was not able to trace completely because of its direction in the fossilization. Parawonnacottella UEDA, 1996

Diagnosis. Anterior margin smooth, where the nodal line reaches Parawonnacottella signifer n. sp. the margin; cell between Rs and M1 almost equally separated by (Figs. 2, 6B) the nodal line: CuA1 aligned with stem of CuA; CuA2 strongly curved towards CuP just after the branch with CuA1. Diagnosis. Wing markings and bands distinct. Name. After wing markings and bands present. Parawonnacottella araripensis UEDA, 1996 (Figs. 1, 6A) Description. Species known from single right fore wing, basally cut artificially. Antero-basal region of fore wing partly missing; Head small, 1/3 of the width of mesothorax; eyes small anterior margin of irregularly sinuous; other veins arranged as 24 Kyoichiro UEDA

(CP) nodal line

(ScP)

AP AA R (RA) CuP CuA CuA M M M 4 3 M R 2 (MP 2 (MP 1 S (MP 1 (RP) (MA) 3/4 2 ) 1 ) )

Fig. 1. Parawonnacottella araripensis Ueda. Dorsal, complete adult specimen. those of Parawonnacottella; wing was wrinkled near distal margin Description. Species known from single left fore wing; most below Rs. of CP, basal portions of Rs, CuA1 and CuA2, apical portions of

Wing markings; six dark grey patches beyond ScP present Rs, M1-4 and CuA1, and CuP are missing. Wing triangular with in the cell below costa and R; dark grey band broad, running apical region pointed; costa rather straight and not so indented at from R to M3 beyond nodal line, separated into two faint and nodus; CP only discernable at middle; ScP missing just from R irregular bands below M3 and reached to inner margin; pale but distinct to Costa; cell between Rs and M1 narrow and distal grey submarginal patch from R to M3 and apical patch faint region beyond nordal line 1/2 as long as basal region. R, Rs and and irregular; discal region from base of M pale grey basally; M divided trichotomously from antero-basal edge of discal cell; postnodal region pale grey; four small patches discernable in cell stem of M distinct; M-CuA indistinct but only traceable from below CuP. M; M1-2 branched dichotomously but stem of M3-4 very short and almost from nodal line. Cross vein r-m short. CuA branched

Holotype. Right fore wing. KMNH IP 000,007 dichotomously; CuA1 evenly curved anteriorly; CuA2 strongly Dimension. Length 52 mm (from the cut-off basal margin to curved apically toward tornus; CuP untraceable; AA and AP apex), width 28 mm. almost parallel and straight to inner margin.

Remarks. Grey bands and markings are well preserved. This is Holotype. Left fore wing. KMNH IP 000,008 the third species of Parawonnacottella and the first record with Dimension. Length 63 mm, width 23 mm. distinct wing markings. Remarks. This specimen is similar to Colossocossus rugosa MEnOn et al., but apparently different from it by the following Genus Colossocossus MENON, HEADS & MARTILL, 2005 character-states: Colossocossus acutus n. sp. 1) costa not so indented at the nodus, (Figs. 3, 6C) 2) R, Rs and M from antero-basal edge of discal cell, 3) fore wing length longer than rugosa (57 mm). Diagnosis. Wing length 63 mm. Nodus not so indented. R, Rs, M divided from the antero-basal edge of discal cell. Unfortunately the genus Colossocossus was established Name. After apical region of the wing pointed. based on rather poor specimens (MEnOn et al., 2005), i. e., the basal wing regions of the species in this genus are not well preserved. The stem of M (crossvein of MEnOn & HEADS, 2005) New and additional palaeontinid species from the Lower Cretaceous of Brazil 25

R (RA) Rs (RP)

M1 (MA)

M2 (MP1)

nodal line M3 (MP2) (ScP)

M4 (MP3/4)

CuA1 (CP) ?

CuA2

M stem CuP AA AP

Fig. 2. Parawonnacottella signifer n. sp. Holotype, right fore wing. was not found in C. loveridgei and C. rugosa (l. c., Figs. 2A-B, 3), from the nodal line (Sc present) and Rs and M not bifurcated but it was observed in C. bechlyi (l. c., Fig. 4), and the generic in C. bechlyi. These conditions clearly suggest that the generic diagnosis was changed from “ triangular antenodal discal region diagnosis of Colossocossus should be revised at least on its basal reaching wing base” to “ reaching wing base or with proximal wing region. The wrinkled texture on the posterior margin of end close to wing base, free of crossveins or with one crossvein” the fore wing is probably the natural condition of large winged (MEnOn & HEADS, 2005: 7). The basal branching patterns on Palaeontinidae (see Parawonnacottella signifer). R, Rs and M are different on each species, i. e., Rs and M are So, here I tentatively placed this new species into the genus bifurcated in C. loveridgei (the basal region of R indistinct), R Colossocossus mainly from the shape of postnodal discal region and M bifurcated then R and Rs bifurcated in C. rugosa, and R and the arrangement of M3-4. 26 Kyoichiro UEDA

R (RA)

Rs (RP)

M1 (MA)

(ScP) M2 (MP1) nodal line

(CP) M3 (MP2)

M stem M4 (MP3/4) M4-CuA

CuA1

CuA2

postnodal discal region CuP AA AP

Fig. 3. Colossocossus acutus n. sp. Holotype, left fore wing.

below CP; distal region of cell between Rs and M beyond nodal line shorter than basal region; cross vein r-m short; M branched Genus Baeocossus MENON, HEADS & MARTILL, 2005 dichotomously; M3 and M4 abruptly branched at antero-distal Baeocossus rotundatus n. sp. edge of postnodal discal region; CuA branched dichotomously;

(Figs. 4, 6D) CuA1 slightly shifted posteriorly but almost aligned with stem of

CuA; CuA2 moderately curved towards CuP after postero-distal Diagnosis. Small species; fore wing length 26 mm with round edge of postnodal discal region; CuP, AA and AP almost straight apex; M3 and M4 branched at antero-distal edge of postnodal to inner margin. discal region. Name. After round apex of fore wing. Holotype. Almost complete specimen, KMNH IP 000,009 Dimension. Fore wing length 26 mm, width 10 mm. Pronotum Description. Almost complete specimen in dorsal aspect; the 11 mm wide. basal region of both fore wings slightly missing; hind wings present; each vein of hind wing not clearly traced; terminal Remarks. This new species differs from the known Baeocossus region of abdomen missing; wing markings and bands present species by the following characher-states: but indistinct. 1) small size (fore wing length 26 mm), Fore wing narrow, triangular with round apex; nodus slightly 2) apex of fore wing round, indented; CP present and parallel with costa to nodal line; R(RA) 3) M3 and M4 abruptly branched at antero-distal edge of and Rs(RP) from the base of wing; R(RA) below Rs(RP) near the postnodal discal region. base but not anastomosing each other and running almost parallel 4) markings and bands present below costa and on whole fore New and additional palaeontinid species from the Lower Cretaceous of Brazil 27

M stem

R(RA) (CP) Rs(RP)

nodal line (ScP)

AP AA R (RA)

CuP CuA CuA M M 4 (MP 1 2 3 M (MP Rs (RP) M 2 (MP 3/4 1 (MA) ) 2 ) 1 )

Fig. 4. Baeocossus rotundatus n. sp. Holotype, dorsal adult specimen.

wings. branching point of M1-2; M3 and M4 bifurcated; M3 from antero-

In addition to above character-states, the separated R (RA) distal corner of antenodal discal region; M4-CuA cross vein and Rs (RP) from the base to the stem of M is plesiomorphic distinct and from a little posterior to branching point of M3-4; but good diagnostic character for the genus Baeocossus. MEnOn CuA bifurcated; CuA1 aligned with stem of CuA; CuA2 evenly & HEADS (2005) considered R(RA) as Sc and also showed this curved towards CuP and almost straight to distal margin; clavus condition in the fore wing of B. fortunatus (paratype, Fig. 3), but with two anal veins (AA and AP) from base. Wing markings and no traces of them in the holotype (Fig. 2). bands well preserved; scattered markings present on basal region of fore wing, especially cells below costa and Rs (RP); six dark Baeocossus muratai n. sp. grey bands traceable; 1st from costa below anastomosing point of (Figs. 5, 6E) R (RA), Rs (RP) and M, to CuP distinct; 2nd broad, from costa to discal cell, then along nodal line basally to inner margin; 3rd

Diagnosis. Small species; fore wing length 28 mm with round from costa around ScP to M2; fourth broad, from costa through apex; wing markings and many minute pits well preserved. cross vein r-m and irregularly divided into two below M3; basal Name. After the donator Mr. Masayuki MURATA, who kindly one along cross vein m-cu to inner margin; distal one interrupted donated this important and beautiful specimen to KMNH. by each vein and reached to tornus; fifth post medial, from

costa, with indistinct white region below Rs to M3 and gradually

Description. Almost complete adult specimen in dorsal aspect; terminating to M4; sixth subterminal, shifted distally below R head broken; no traces of legs; thorax broken, 8 mm long, 10 mm (RA) to M1; a series of indistinct shades at each vein on distal wide; lateral margin of pronotum pointed; abdomen 6 mm long margin. and segments indistinct; total body length 14 mm. Hind wing small, 16 mm long and 9 mm wide, with round

Fore wing, 28 mm long, 13 mm wide, and triangular with apex; Rs and M1 bifurcated beyond mid wing; M2 and M3 from round apex; costa indented at nodus; CP complete from base to base gently curved to margin; CuA1-A2 bifurcated at base; CuP ScP; R (RA) from base and slightly anastomosing with Rs (RP) present. Broad bands from costa to margin, shifted distally at and M stem at antero-basal corner of antenodal discal region; R each vein, and running on middle and subterminal regions; a (RA), Rs (RP) and M stem running separately to nodal line; cell large distal patch below Rs. between R (RA) and M stem narrow; distal region beyond nordal line shorter than basal region; M-CuA cross vein slender but Holotype. Almost complete specimen. KMNH IP 000,006 distinct; M branched dichotomously; M1 strongly angled to Rs Dimension. Fore wing length 28 mm, width 13 mm; hind wing from branching point; cross vein r-m short and just distad from length 16 mm, width 9 mm; pronotum length 8 mm, width 10 28 Kyoichiro UEDA Rs M 1 M 2 M 3 CuA 1 CuA CuP 2 M stem R(RA) (CP) AP Rs (RP) M-CuA nodal line AA (ScP) CuP CuA 2 CuA 1 M 4 (MP M 3 M (MP R(RA) 3/4 2 (MP M ) 2 1 ) Rs (RP) (MA) 1 )

Fig. 5. Baeocossus muratai n. sp. Holotype, dorsal complete adult specimen. New and additional palaeontinid species from the Lower Cretaceous of Brazil 29

A B

C D

E

Fig. 6. A: Parawonnacottella araripensis Ueda, B: Parawonnacottella signifer n. sp., holotype, C: Colossocossus acutus n. sp., holotype, D: Baeocossus rotundatus n. sp., holotype, E: Baeocossus muratai n. sp., holotype. 30 Kyoichiro UEDA

Fig. 7. Fossil localities of Araripe. A: One of outcrops. Photo by BRITO, B: One of fossil insects sites, quarry of Nova Olinda. Photo by Annesuse RAqUET-SChWICKERT.

Table 1. Palaeontinid genera from the Cretaceous (modified from MEnOn, HEADS & MARTILL, 2005). Liaocossus (MEnOn et al., assigned it as a synonym of Ilerdocossus) and Miracossus from the of NE are excluded from this table based on the age of the locality as indicated Late by REn et al. (1998) Age Taxon Locality and horizon Reference

L. Aptian Parawonnacottella Nova Olinda, Ceará, Brazil: Cato Fm UEDA (1997) (120MA) Catocossus same above MARTInS-NETO (1998) Baeocossus same above MEnOn et al. (2005) Colossocossus same above MEnOn et al. (2005)

Hauterivian Cicadomorpha Daya, Chita, Transbaikalia, Siberia: ShChERBAKOV (1988) (132MA) Glushkovskaya Fm

L. Berriasian~ Pachypsyche Sierra del Montsec, Lérida, : MEUnIER (1902) E. Valanginian Caliza con Caraceas Fm (140MA) Montsecocossus same above GOmEz-PALLEROLA (1984) Ilerdocossus (syn. sama above GOmEz-PALLEROLA (1984) Wonnacottella) New and additional palaeontinid species from the Lower Cretaceous of Brazil 31 mm; abdomen length 6 mm; total body length 14 mm. gave useful comments. I am indebted to Prof. DOnG Ren, College of Life Science, Capital Normal University, Beijing and Dr. Remarks. This beautiful specimen has been featured as cover Mamoru OWADA, National Science Museum, Tokyo, for literature photo and on colour plate 12a in the recent book “ The Crato reseach. Dr. Paulo M. BRITO, Departamento de Biologia Fossil Beds of Brazil - Window into an Ancient World ” by e Vegetal, Universidade do Estado do Rio de Janeiro and Mrs. MARTILL et al. (2007), and was tentatively determined as Annesuse RAqUET-SChWICKERT, Sulzbachtal, allowed me to use Baeocossus cf. fortunatus in this book. It has a similar wing their photos of the localities. venation like Baeocossus fortunatus MEnOn & HEADS, but differs REFERENCES from it by the following character-states: 1) small size species, body length 14 mm, fore wing length 28 DWORAKOWSKA, I. 1988. Main veins of the wings of Auchenorrhyncha mm and hind wing length 16 mm, but these are 17 mm, 35 (Insecta, Rhychota: Homoptera). Entomologische Abhandlungen 52 mm and 15 mm in fortunatus, (1) 3: 63-108. 2) wing markings and bands clearly different from fortunatus, KUKALOVá-PECK, J. 1991. Fossil history and the evolution of hexapod which has no distinct bands, structures. in NAUmAnn, I. D. et al. (eds.). The insects of 3) apices of both wings are round, while pointed in frotunatus. , a textbook for students and research workers. Vol. 1 (2nd edition): 141-179, Melbourne University Press, Melbourne. On B. finchae MEnOn, HEADS & MARTILL, the type species MARTILL, D. M., BEChLY, G. & R. F. LOVERRIDGE (eds.). The Crato of the genus Baeocossus is rather poorly preserved and no good Fossil Beds of Brazil. Window into an Ancient World. xvi + 625 character-states mentioned, but it is also large species with fore pp, Cambridge University Press, Cambridge. wing length 31-35 mm long (MEnOn & HEADS, 2005: 3). MEnOn, F., S. W. HEADS & D. M. MARTILL, 2005. New Palaeontinidae (Insecta: Cicadomorpha) from the Lower Cretaceous Crato The fore wing venation was completely preserved in this Formation of Brazil. Cretaceous Research 26 (2005): 837-844. species and one of good examples to discuss the venation system MEnOn, F. & S. W. HEADS, 2005. New species of Palaeontinidae of Palaeontinidae in future. Especially, R (RA) and Rs (RP) (Insecta: Cicadomorpha) from the Lower Cretaceous Crato are complete from the base and running separated each other Fromation of Brazil. Stuttgarter Beiträge zur Naturkunde. Series except for one anastomosing point with the stem of M at basal B (357): 1-11, 6 Figs., 1 Tab. 1/3 of discal cell. On the hind wing I considered that the anterior UEDA, K., 1997. A new Palaeontinid species from the Lower margin of hind wing was covered by the inner margin of fore Cretaceous of Brazil (Homoptera: Palaeontinidae). Bulletin of wing and the R (RA) was not traceable on this specimen. Kitakyushu Museum of natural History, 16: 99-104. WhALLEY, P. E. S. & E. A. JARzEmBOWSKI, 1985. Fossil insects from ACKNOWLEDGEMENTS the lithographic limestone of Montsech (late Jurassic-early Mr. Masayuki MURATA, Director of the Kiyomizu Cretaceous), Lérida Province, Spain. Bulletin of the British Sannenzaka Museum, Kyoto kindly donated the important Museum (Natural History), London, (Geology series), 38 (5): species to KMNH. Dr. Günter BEChLY, Staatliches Museum für 381-412. Naturkunde, Stuttgart reviewed an early draft of this paper and