ACTA GEOLOGICA HISPANICA, v. 35 (2000), nº 1-2, p. 165-169

New cantacaderid lace bugs from Dominican amber (Heteroptera: , Cantacaderinae)

V.B. GOLUB(1) and Y.A. POPOV(2)

(1) Voronezh University, Universitetskaya pl., Voronezh, 394693, Russia. (2) Paleontological Institute RAS, Profsoyuznaya str. 123, Moscow 117686, Russia. E-mail: [email protected]

AB S T R A CT

Ne w fossil tingids, representatives of the small subfam i l y Cantacaderinae (Tingidae) from the Oligocene Dominican amber, are describ- ed and discussed. The fossil species Eocader baby r u s s u s n. sp. belongs to the recent neotropical genus Eo c a d e r of the tribe Phatnomini.

Keywo rd s : Heteroptera. Tingidae. Cantacaderinae. Phatnomini. Eocader baby r u s s u s n.sp. Dominican amber.

IN T RO D U C T I O N Ne w fossil species of the peculiar cantacaderid lace bugs were preserved in amber from the Dominican Repub- Fossil tingids are quite rare among other known fossil lic. This amber varies in age from Lower Miocene to Mid bugs, especially they are ver y few in Mesozoic heteropte- Oligocene, i.e. 23-30 million years old (Grimaldi, 1995). ran faunas (Pop o v, 1989; Golub and Pop o v, 1998a; Golub We support the conclusion made by Iturralde and MacPhee and Pop o v, 1999). Most fossil lace bugs described and (1996) that the age of Dominican amber in the Dominican mentioned in publications from Cenozoic Dominican and Re p u b lic is most probably of the late Early Miocene - early Baltic ambers mainly belong to the plesiomorphic subfa- Middle Miocene (15 to 20 million years old). The present mi l y Cantacaderinae (especially in the Baltic amber) and paper deals with the first tingid description from Domini- are represented almost exc e p t i o n a l l y by the tribe Phatno- can amber (all amberiferous sites are unfortu n a t e l y un- mini (Golub and Pop o v, 1999). Some of them, e.g. Si n a l - kn o wn) based on 4 specimens (2 males and 2 females) whi l e da baltica (DRAKE), as well as species of the genus Pa- some other undescribed tingids from Dominican amber be- le o c a d e r GOLUB AND POPOV 1998, are quite common. long to the other subfam i l y and to the recent Am e - rican genus Ga r gap h i a (Golub and Pop o v, 1999). During the visit of one of the authors (Yu. P .) to the Smithsonian Institution (Washington) and the Am e r i c a n Museum of Natural History (New York) in 1994 and also SY S T E M A TICS due to continuous effo r ts by Dr.W. W eitschat (Hambur g Un i versity) three ver y fine collections of the Dominican Ord e r : He m i p t e r a amber inclusions have been studied and the tingids from Su b o rd e r : He t e r o p t e r a them were sorted for determination and description. In f ra o rd e r : Ci m i c o m o r pha

165 Fam i ly : Tingidae LAPORTE, 1832 with 1-2 finest cells; median carina stretching from pos- Su b f a m i ly : Cantacaderinae STAL, 1873 terior margin of calloused convex area to posterior margi n Tri b e : Phatnomini DRAKE AND DADIS, 1960 of pronotum. Pronotal disc densely punctuate (except cal- loused small area); anterior margin distinctly emargi n a t e , GENUS Eo c a d e r DRAKE AND HAMBLETON 1934 el e vated in form of rudimental vesicula with 1 transver s a l ro w of small cells. Paranotum in anterior part consid- Eocader babyr u s s u s GOLUB AND POPOV n.sp. er a b ly widened, with 3 row of cells (3rd intermediate row Figure 1 represented by a single cell only) narro wing backwar d and with 1 row of rather small cells along most part of Type species: Eocader vergra n d i s DRAKE AN D their length; ext e r nal edges in anterior third emargi n a t e ; HA M B L E T ON 1934; recent species. posterior margin of pronotum almost straight and slightly waved , without triangular projection. Scutellum comple- Type material: Holotype male (submacropterous te l y exp o s e d , trapezoidal form. fo r m); label “Amber: Dominican Republic, Oligo-Mio- cene, specific provenance unknown. Purched in Santo Submacropterous form. Hemelytra surpassing far be- Domingo by D.Grimaldi, AMNH DR-8-383.”; wel l yond top of abdomen, sharpl y narro wed at base. Costal pr e s e r ved (Fig. 1.1) and deposited in the American Mu- area rather wide, practically along whole length with 2 seum Natural History, New Yor k . ro ws of moderately large cells, mainly quadrangular-p e n - tagonal form. Subcostal area inclined (this area of right De s c r i p t i o n : Holotype, male. Small, less than 2 mm he m e l ytron of holotype slightly flattened), with 4 rows of long. Body oblong, bare and general colour brown (pro- cells along most length and with 4 transversal veins eleva- ba b ly originally yel l ow - gr ay ) . ting more than other veins. Discoidal area in broadest pla- ce with 4 rows of rounded, irre gular form cells, and with Head long, 1.7 times as long as wide (from clypeus tip 2-3 transversal strongly elevated veins. Sutural area at the to hind margin of eyes) and 1.34 times as long as wide ver y base of hemelytra with one row of cells at the level of (from clypeus top to anterior emarginate margin of prono- anterior angle of discoidal area, slightly broadening and at tum). Surface of head fine punctate. Vert e x between eyes this place with a single cell of 2nd row; further backwar d s and occiput depressed, considerably lower than convex along discoidal area with 1 row of cells and near by apex frons; this depression runs obli q u e l y outside in form of of widened sutural area 5 rows of cells; tips of sutural are- sulcus in front of each eye fusing backwards and Y-l i ke . as (membranae) of both hemelytra almost fully over l a p - Ey es convex, rising above head surface, directed upwar d s ping in repose. Clavus distinctly separated by sutura from and to the side. Head with 5 denticles: unpaired clyp e a l corium, in the broadest place with 4 oblique rows of cells; and paired frontal and jugal ones. All denticles located far exterior row is separated at the level of vein from others. from eyes, especially clypeal and jugal once; clypeal den- Hypocostal plate (lamina) with one row of cells. ticle conical with thick base, almost equal as wide as the base of clypeus, directed obli q u e l y upwards and forwar d s Scent gland openings located by apex of anterior-e x- and covering (overlain from above) almost 2/3 length of terior angle of metapleurit, slightly elevated and broad- cl ypeus; frontal denticles conical with thick base and ened at this place. Rostrum long, surpassing backwar d s smoothed top, directed obli q u e l y forwards, upwards and anterior margin of fifth abdominal sterni t e . to sides; jugal denticles ver y short, rather thin and wea k l y cu r ved , with sharp apex, located lower and in front of cly- Dimensions (in mm): Body length to the tip of hem- peal one, directly above upper edge of bucculae. An t e n - el ytra 1.86, width 0.87; length of pronotum 0.37, width nae long and thin; 1-2 joints ver y short, 2nd joint far from 0.57; length of head from clypeal tip to hind margin of eyes reaching clypeal top; 3rd joint thinnest and longest. An - 0.34, to anterior margin of vesicula (along middle line) te n n i f e r us tubercles ver y small and almost equal in length 0.39, width 0.29; width of vert e x 0.11; ratio of antenno- to 2nd joint. Bucculae anteriorly opened, hardly protru- meres I-IV as 0.05:0.045:0.64:0.16. ding forward beyond apex of clypeus; anterior half with 2 horizontal rows of cells, posterior one has 1 row of cells. Var i ab i l i t y : Judging by the preservation to diffe r e n t de grees of body parts of paratypes (1 male and 2 females) Pronotum 1.54 times as wide as long; pronotal disc cl ypeal denticle may be shorter and overlap one third - co n vex, with 3 longitudinal low carinae lateral ones of one half of the length of clypeus (Fig. 1.2), broadened them wea k l y protruding; median carina in its middle part front parts of paranota can be only with two rows of cells,

166 Figure 1. 1.- Habitus of Eocader babyrussus n. sp.; 2.- Drawing of the holotype, AMNH DR-8-383. at least females can be brachypterous (female paratype) atype with preserved antennae the length of the third and with fusing but not overlapping sutural areas of hemely- fo u r th joints are corre s p o n d i n g l y 0.53 and 0.16 mm; a tra along the whole of their length, and subcostal and dis- si g n i f icant difference in the length of the third joint of an- coidal areas of hemelytra in their broadest part can be tennae in males and females of the same species is a reg- with five rows of cells (at least in females). The length of ular phenomenon in fam i l y Tingidae. Besides the above, the third and fourth antennal joints in paratype of male frontal denticles of head can be more rounded and flat on ranges corre s p o n d i n g l y 0.65 and 0.14 mm. In female par- the top as it is shown in figure 1.2.

167 Type material: Paratype ? male; amber Dominican straight hind margin. The differences in the structure of Re p u b lic; insect strongly damaged (Fig. 1.2) and deposi- pronotum of E. vergra n d i s and E. babyrussus n.sp. are ted in the collection G. Herrling (NW Germ a ny ) , on l y of a special character. In the same way, the presence In v.N r .DB W2. Paratype female; label “12533 - amber or absence of any pair of head denticles cannot serve as a Dominican Republic, C. Drake collection”; insect signifi- su f ficient ground for distinguishing this new species as an ce n t l y damaged and deposited in the National Museum of independent genus. In the composition of the modern ge- Natural History, Smithsonian Institution. Paratype fema- nera of Tingidae there are often species both with carinae le; label “125445 - amber Dominican Republic, C. Drake and without them, e.g. in Galeatus inermis the head’s ca- collection”; insect strongly damaged and deposited in the rinae are absent altoge t h e r , while in other species of this National Museum of Natural History, Smithsonian Insti- genus there are five long carinae. tu t i o n . Oligo-Miocene E. babyrussus n.sp. is very close to Et y m o l o gy: The name of the species reflects a curious the Eocene genus Intercader GOLUB AND POPOV similarity in the form of head with the quite fam o u s 1998b, from Baltic amber. The only species of this genus Oriental pig Ba byrussa baby r u s s a . I. weitschati similar to E. babyrussus n.sp. has a depres- sed vertex between eyes and five head denticles inclu- ding the jugal one. The main differences between species DISCUSSION I.weitschati are the presence of five pronotal carinae and the absence of a sharp narrowing of hemelytra at their The absence of stenocostal area of hemelytra and the base. The paired short lateral carinae closely positioned presence of median clypeal protrusion are the features of on each side of pronotum in Intercader can be looked Phatnomini. The depression of vert e x between eyes with upon as a result of the break of initial carina and diver- its continuation in the form of two oblique gro o ves in gence of its fragments in the process of evolution. In any front of eyes (Y-shaped form) and hemelytron sharpl y case the difference in the number of pronotal lateral ca- n a rr owing at the base - all these are the characteristic fe- rinae is far from being always considered in systematics atures of the small Neotropical recent genus Eo c a d e r of Tingidae as generic difference, for instance in genus (D r a k e and Hambleton, 1934). Judging by the diagnosis Derephysia. The differences in the form of hemelytra ba- of genera in the excellent work of Richard C. Fro e s c h n e r se are probably more significant because of the unique- (1996), these specific features of Eo c a d e r are unique ness of this feature in Miocene and modern species of among Phatnomini. The broadened front parts of parano- genus Eocader. However the depression of the vertex in tum and their emarginated exterior edges are the features Eocader and Intercader is also a rare feature among Tin- that bring together the fossil Eocader baby r u s s u s n. s p . gidae, whose independent appearance in phylogenetic with the type species of the genus E. vergra n d i s DR A K E groups seems to be less possible in comparison with re- AND HAMBLETON found in Brazil. lated groups. The discovery of the modern genus Eoca- der in Dominican amber and the closest fossil genus In- The main difference between the Miocene species and tercader in Baltic amber, on the one hand, points to it the modern E. vergra n d i s is the absence in the latter of ju- (Eocader) being rather old (beginning with Early Ceno- gal denticles. Besides, the frontal and clypeal denticles in zoic period) and, on the other hand, it can be compared E. babyrussus n.sp. are located closer to the top of head to the recent species of this genus. In the Tertiary period as compared to E. vergra n d i s . In addition to well exp r e s - there existed significantly closer zoogeographical links sed medial carina, the holotype and paratypes of the spe- between the faunas of Eurasia and South America, as cies described here have ext r e m e l y low lateral carinae on compared to the present time (Golub and Popov, 1998 a). pronotum. In brachypterous specimens of E. vergra n d i s , pronotum has one carina and emarginate hind margin, in the macropterous ones - with three carinae and emargi n a - ACK N OW L E D G M E N T S te hind margin (Fro e s c h n e r , 1996). In the submacropte- rous holotype E. baby r u s s u s n.sp. pronotum has a wel l We are extremely grateful to Prof. Richard. C. Froeschner exp r e s s e d , although ver y low, medial carina and slightly (NMNH, Washington), Dr. David A. Grimaldi (AMNH, New rising lateral carinae, while the hind margin of pronotum York) and Dr.Wolfgang Weitschat (GPIM, Hamburg) and Mes- is practically straight. At least one of the paratypes of the sers G. Herrling (Bramsche, NW-Germany). We are thankful to described species is a brachypterous female. Its pronotum the reviewers Dr. E. Guilbert and another anonimous one who has also three carinae (lateral ones are ver y low) and a provided a critique of the manuscript and helped in preparing

168 the final text. We are also especially indebted to Olga Seliva- Golub, V.B , Pop o v, Yu.A., 1998b. Cantacaderid lace bugs from the nova (Voronezh University, Russia) for excellently fulfilled ori- Baltic amber (Heteroptera: Tingidae, Cantacaderinae). Mitt. ginal figure 1.2. Ge o l . - P allllaont.Inst. Univ. Hambur g, 81, 223-250. Golub, V.B ., Pop o v, Yu.A., 1999. Composition and evolution of Cretaceous and Cenozoic faunas of bugs of the superfam i l y REFERENCES Tingoidea (Heteroptera: ). Entomol. Prob- lems (in press) Dr a k e, C.J., Hambleton, E.J., 1934. Brazilian Tingidae (Hemip- Grimaldi, D., 1995. On the age of Dominican amber. In: K.B. tera), Par t I. Rev.Ent., Rio de Janeiro, 4, 435-251. Anderson and J.C. Crelling (eds.) Am b e r , resinites, and fos- Fro e s c h n e r , R.C., 1996. Lace bug genera of the Wor l d , I: Intro- sil resines. Amber and Resinites, Chemical Society Sympo- duction, Subfam i l y Cantacaderinae (Heteroptera: Tin g i d a e ) . sium volume (Washington, D.C., Aug.1994), 1-11. Smithsonian Contrib.Zool., 574, 1-43. It u r ralde, M.A., MacPhee, R.D.E., 1996. Age and paleoge o - Golub, V.B ., Pop o v, Yu.A., 1998a. Cretaceous and Pal e og e n i c graphical origin of Dominican amber. Science, 273, 1850- faunas of bugs of the superfam i l y Tingoidea of the eastern 18 5 2 . and wes t e r n Hemispheres, their relationships and evol u t i o n . Pop o v, Yu.A., 1989. New fossil (Heteroptera+Cole- First Pal e o e n t o m o l o gical Conference, Moscow, 1998 (Abs- o rr hyncha) from the Mesozoic of Mongolia. N. Jb. Geol. Pa- tracts), 7. laont. Mh., 3, 66-181.

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