POLSKIE PISMO ENTOMOLOGICZNE P O L I S H J O U R N A L O F E N T O M O L O G Y

VOL. 69:143-153 Gdynia 31 July 2000

First (Heteroptera: ) from Baltic amber ALEKSANDER HERCZEK

Department ofZoology, Silesian University, Bankowa 9,40-007 Katowice, Poland e-mail: herczek~us.edu.pl

ABSTRACT. Fossil Miridae ofthe subfamily Phylinae from Baltic amber are reported for the first time. A new genus Hallodapomimus gen. n. including two new species H. elektrinus sp. n. and H. succinus sp. n. are described and illustrated. The new genus is included in the tribe Hallodapini.

KEY WORDS: Heteroptera, Miridae, Phylinae, Hallodapini, Baltic amber, fossil plant bugs.

INTRODUCTION

Most reports on heteropteran amber inclusions of the family Miridae concern representatives of the subfamilies Isometopinae and Cylapinae (CARVALHO 1966, 1985, CARVALHO & POPOV 1984, GERMAR & BERENDT 1856, JORDAN 1944, HERCZEK 1991a, 1993, HERCZEK & POPOV 1997a, b, PoPov & HERCZEK 1993). Species which belong to other mirid subfamilies have seldom been described (HERCZEK 1991b, HERCZEK & GORCZYCA 1991, HERCZEK & POPOV 1992), and there are no data on the representatives of Phylinae (in spite of available inclusions), even though this group has been studied extensively (e.g. by SCHUH 1974, 1984). This shortage seems to be resulted from the large number of extant genera, which constitute the latter subfamily (about 300), and to the wide distribution of their representatives, which occur in tropical, subtropical and temperate zones (ScHuH & SLATER 1995). The situation is firther complicated by the fact that in the analysis there are important characters ofgenitalia, which makes a comparison of fossil and recent forms difficult. The Phylinae, like Mirinae and Orthotylinae are rare among amber inclusions. The species described represent the tribe Hallodapini. The tribe sometimes is included in the subfamily Hallodapinae, sensu WAGNER 1973. Since both sexes ofthe same species have been found in amber, it has been established that dimorphism, frequent in recent Miridae, occurs also in this group. Moreover, certain characters have been observed 144 Polskie Pismo Entomologiczne 69 (2) which make bugs similar to ants (e.g. the basal part of abdomen narrowed), which again is typical of many recent representatives of Hallodapini, Leucopterophorini, Auricillocorini, Pilophoririi.

Acknowledgements I would like to thank Dr. Wolfgang Weitschat and Mr. Jens Holt for the loan of amber inclusions. I also thank Mr. Jacek Serafin who kindly helped me to prepare the amber for study and also for the loan of specimens. Special thanks are due to Dr. Yu.A. Popov for his valuable suggestions and improvements ofthe manuscript.

SYSTEMATIC PART Genus: Hallodapomimus HERCZEK, 1998 Type species: Hallodapomimus elektrinus sp. n. by present designation.

Diagnosis The representatives of the genus Iallodapomimus differ from other Baltic amber Miridae in the shape of pronotum, the width of pronotal collar and the pattern of pale patches on hemelytra. The arcuate incision of the lateral margins of hemelytra, make bugs resemble ants. Derivatio nominis The generic name has been coined from the name ofthe recent genus Hallodapus and a Latin word mimus, which refers to similarity. Description Body length up to 6.5 mm, lateral margins of hemelytra slightly notched in their mid part. Body almost smooth, covered with sparse, short, clinging hairs. Hemelytra with two pale yellow patches, one close to the wing base reaches mid part of clavus, the other one in the basal part of cuneus. Head subvertical to subhorizontal, convex in the hind part. Eyes relatively big reaching half height of head and almost touching the anterior margin of pronotum. Vertex slightly beneath the upper margin of eyes. Foveae antennalis situated below 2/3 eye height, almost touching the inner margin ofeye. Antennae covered with very short, clinging setae, first antennal segment slightly longer than head, second segment almost cylindrical or club-shaped. Rostrum relatively short, reaching coxae of hind legs, first rostral segment touching the anterior margin of prostemum. Other segments equal in length. Pronotum 1.5 times broader than long in basal part, in the apical part strongly narrowed, collar-like. Latero-posterior angles rounded, slightly bevelled and flattened. Calli distinct, moderately convex. Collar distinct, flattened. Mesoscutum moderately exposed, scutellum slightly wider than long, markedly convex, separated from mesoscutum by a transverse dent. Hemelytra with radial and claval veins very weakly marked. Cuneus HERCZEK A.: First Phylinae (Heteroptera: Miridae) from Baltic amber 145 relatively long, about 1.5 times longer than broad. Membrane hazy, delicately creased along the long axis, with two distinct, closed cells (the smaller one narrow and elongated). All femora slender, elongated; hind femora extending slightly beyond abdomen. Tibiae with two rows ofspines on inner surface; length ofspines less than diameter oftibiae. First tarsal segment swollen, almost twice as long as the next one and subequal in length to third tarsal segment. Claws thicker at base, moderately crooked; parempodia hair-like, parallel; pulvilli very small.

Hallodapomimus elektfinus sp. n.

Hallodapomimus elektrinus: HERCZEK 1998, nom. nud.

Description Male. Macropterous form. Body elongated. Body ground colour dark brown. Hemelytra with two creamy patches: one close to the basal part ofwing covers corium, the mid part of clavus and reaches commissura clavalis, the other occupies half ofthe basal part of cuneus. Apical part ofscutellum pale. Dorsal surface smooth, impunctate, covered with sparse, very short and clinging setae. Head subvertical, almost twice as broad as long, clypeus distinct, protruding above frons. Genal cones distinct. Eyes relatively large, slightly protruding sidewards, somewhat elevated above the vertex surface and slightly removed from the anterior margin of pronotun. Antennae inserted above the lower margin of eye, at 2/3 its height. Fovea antennalis touches the inner margin of eye. Second antennal segment almost cylindrical (only slightly expanded apically). Third segment almost twice as long as the fourth one; both thinner than the second. Rostrum relatively short, reaching hind coxae. All rostral segments (except the first one, which is somewhat shorter) are equal in length. Pronotum tapers strongly at 1/3 its length and forms a collar-like structure. Collar distinct, broad and flat. Calli distinct, slightly convex, separated in the middle, reaching the lateral margins of pronotum. Mesoscutum moderately exposed. Scutellum convex, more than twice as long as mesoscutum. Commissura clavale slightly longer than 1.5 length- of scutellum. Hemelytra with the lateral margins almost parallel, slightly concave at 1/3 their length (the widest part at a level with cuneal fracture). Membrane hazy, delicately creased along the long axis, with two closed cells: the smaller one narrow and elongated, the other with the posterior proximal angle rounded. Abdomen relatively narrow, slightly tapering in the basal part. All femora slender, elongated, hind femora with short spines scattered on the inner surface. All tarsi three-segmented: first tarsal segment slightly thicker than others, almost twice as long as the second one and subequal in length to third tarsal segment. Claws thicker at base, moderately crooked; parempodia parallel, hair-like; pulvilli very small. Female. Macropterous form. Body length 6.2 mm. Body ground colour brown. Hemelytra and the apical part ofscutellum with pale patches. One begins near the basal 146 Polskie Pismo Entomologiczne 69 (2)

,t

Fig. 1. Hallodapomimus elektrinus sp. n., holotype, male, coil. Museum ofthe Earth, Warsaw.

Fig. 2. Hallodapomimus elektrinus sp. n., paratype, female, Geologisch-Paldontologisches Institute und Museum, Universitdt Hamburg (coll. Jens von Holt). HERCZEK A.: First Phylinae (Heteroptera: Miridae) from Baltic amber 147 wing part ad occpis about 1/3 length of hemnelytra, the mid part of clavus and reaches commissura clavalis, the ote covers the basal half of cuneus. DoMsal surface smooth, cveed, wifth sparse, ort, clinging set&e. Frons, clypeus and first and second rostral sments covered with somewhat longer and denser seta. Head subhorizontal, the Posteior of vetex slightly convex. Diameter of eye smaller than weakly convex vertex, eye occupies somewhat more than half height of head, its posterior margin almost straight In lateral aspect clypeus does not protide above frons. Buccula and genae broad. hnta~ inserted at 2/3 eye height, foveae antennalis touch the inner margin of eye. First antal sem long, longer an had, second segment clib-shaped, third segment about 2.3 tines longer than the fourth one. All segments covered with short, clinging setae. Collar brd flatened, brader th the diameter of first antennal segment. Pronotum tapers in the apical third of its length; this part is covered by flat calli distinctly sparated from the posteror, stongly elongated part of pronotu. Latero-posterior angles bevelle the posterior marg weakly convex, with a shallow notch in the mid part. Mesosctwn covered, the exposed part 5 times shorter than the slightly convex scutellumn. Coriwn weakly dented at sides in te middle of its length. The widest body part slightly above cumeal fare, which forms acnte angl with the long body axis. Membrane with two distict, clsd cells, the bigger almos rectangular. Abdomen tapers basally, extends in the apicl put Hind femora slightly loner than abdomen, covered on the ventral surface with irqp shrt spines, whihh ae absn on fore and mid femora. Tibiae slim, with two rows of shot spies aid covered witfih vy siht, clinging setae. Tarsi with three segments, the diameter of first tarsal segment slighly longer than in the other segments, first segment twice as long as the second one and slightly longer than the third. All tarsal segments covered with shr, proting setae. Claws thicker in the basal part, moderately crooked, with very small pulvilli. Parempodia hair-like, parallel. Male: body length 6.4, width 2.0; length of head 0.68, width 1.3; width of vertex 0.4; width ofeye 0.5, height 0.6; length ofanteal segment: I 0.56, 112.43, III and IV absent; rostal s: 053 11 0.65, 1II 0.65, IV 0.65; length of pronotum 1.2, anterior width 0.74, postrir width 1.55; lngth of n 0.15, width 1.1; length of scutellmn 0.65, width 0.8; claval coaniswe 1m.43; length of hind fenur 2.39, width 0.27, leniet of hind tibia 3.41; length oftarsal segments: 10.34,1 0.18, III 0.31. Female: body ength 6.2, width 1.6; length of head 0.8; width of vertex 0.43; width of eye 0.4, height 0.56; length of segments: 1.1,11 2.85, II 2.04, IV 0.13 (?); rostral segments: 1 0.5, 11 0.62, 1Il 0.62, IV 0.62; length of pronotum 0.99, anterior width 068, posterior width 1.3; length of 0.12, width 0.99 length ofscutellum 0.62, width 0.62; claval caxnmnisuve 1.2; klngth of hind femur 2.79, width 0.25, lenght of hind tibia 3.53; length oftarsal segments: 1 0.36, 110.18, 111 0.31. oype: mal, Balic amber, deposited in the collection of the Museum of the Earth, Wasaw (cel. Y. Poo). 148 Polskie Pismo Entomologiczne 69 (2)

Paratypes: male, Baltic amber, deposited in the collection ofInstitute ofthe Systematics and Evolution ofAnimals PAS, Cracow (coil. Jacek Serafin); male, Baltic amber, deposited in the collection of Geologisch-Paldontologisches Institute und Museum, Universit~t Hamburg (coll. W. Weitschat); female, Baltic amber, deposited in the collection of Geologisch-Paldontologisches Institute und Museum, Universit~t Hamburg (coil. Jens von Holt); male: No. 655, Baltic amber, deposited in the Deutsches Entomologisches Institut in Eberwalde (Berlin, Germany) (coll. C. and H. Hoffeins). Derivatio nominis The name is based on the Greek name ofamber - elektron. Comparison The comparison of the male and female of Hallodapomimus elekirinus shows a considerable sexual dimorphism. The first antennal segment is almost twice longer in females than in males and the second one is distinctly club-shaped in females. The clypeus, genae and the first rostral segment in females are covered with setae (this difference may be attributable to the condition of the preserved specimen) and the mesoscutum is almost totally covered by pronotum, both scutellum and calli are less convex, of equal length and width. In males mesoscutum is distinctly more exposed, while scutellum is convex and markedly longer. There are also differences in the shape of abdomen, which is narrower in males, whereas in females it is considerably expanded apically. The bigger cell on membrane in females is almost right-angled at the posterior proximal edge while in males it is rounded. Similarities of both sexes include the occurrence of short, irregular spines on the inner surface of hind femora, the presence of broad, flattened collar, and a notch in the middle part ofthe posterior margin ofpronotum. The colour pattern is similar as well.

Hallodapomimus succinus sp. n.

Hallodapomimus succinus: HERCZEK 1998, nom. nud.

Description Female. Macropterous form. Body elongated, 5.7 mm long. Body ground colour dark brown. Dorsal surface with pale patches: one covers 1/3 ofhemelytra, it begins at the base ofwing and runs to the mid of clavus, the other covers 2/3 ofcuneus and begins at its base. Scutellum also pale. Dorsum delicately punctate, with very short, clinging setae. Head subvertical, wider than long. Posterior margin ofvertex convex. Eyes developed sidewards, approaching but not touching the anterior margin of pronotum. Their lower, posterior margin (in side view) with a reniform incision. Tylus in lateral aspect protrudes considerably above the plane of frons. Buccula elongated, narrow, gena wide. Antennae inserted at a level ofthe lower inner margin of eye, first antennal segment shorter than the height ofhead, second segment with a distinct club-like swelling at apex, both second and HERCZEK A.: First Phylinae (Heteroptera: Miridae) from Baltic amber 149 third segment covered with very short, clinging setae. Rostrum reaches hind coxae, rostral segments II, III and IV equal in length. First rostral segment longer than the others, reaching almost the middle of xyphus. Collar flattened, narrower than the diameter of first antennal segment. Pronotum tapering at 1/3 its length, convex in the hind part. Calli situated in the narrowed part, flattened. Sides ofpronotum sinusoidally concave, the latero- posterior angles rounded, the posterior margin weakly convex. Mesoscutum moderately exposed, the exposed part almost as long as half length of scutellum. Scutellum weakly convex, equilateral. Lateral margins of hemelytra very weakly notched in the middle, the widest part at the level of cuneal fracture, which is perpendicular to the long body axis. Length and width of cuneus subequal. Cells on membrane distinctly closed. Abdomen does not taper basally. Hind femora overlap the end of abdomen; tarsi with 3 segments, first tarsal segment covered with dense, protruding setae (denser than on segments II and III), thick and almost twice as long as second segment and 1.5 times longer than the third one. Claws crooked, slightly thicker at base. Pulvilli small. Male unknown. Measurements (in mm) Female: body length 5.7, width 1.9; length of head 0.68, width 1.5; width of vertex 0.61; width of eye 0.42, height 0.65; length of antennal segments: I 0.65, II 3.0, III 1.8, IV absent; rostral segments: I 0.76, II 0.65, III 0.65, IV 0.65; length ofpronotum 0.93, anterior width 1.1, posterior width 1.61; length ofmesoscutum 0.27, width 1.14; length of scutellum 0.61, width 0.61; claval commissure 1.14; length of hind femur 2.6, width 0.38, length of tarsal segments (2-pair): I 0.42, II 0.15, III 0.27. Material examined Holotype: female, Baltic amber, deposited in the collection of the Geologisch Paliontologisches Institute und Museum, Universitit Hamburg (Coll. W. Weitschat). Derivatio nominis Named after Latin name ofamber - succinum. Comparison There are distinct differences in morphology of Hallodapomimus elektrinus and H. succinus. The position of the head is subhorizontal (more conspicuous in females) in the former and subvertical in the latter. Also size and shape ofbuccula and genae are different. The compared species differ also in the position of antennae: in H succinus they are inserted almost at the lower margin of eye, while in H. elekirinus at 2/3 of its height. The pronotum in females of H. succinus is shorter and less convex in the hind part, without a notch in the middle of its posterior margin. Also the collar is considerably narrower and the rostrum is longer. The base of abdomen in H. elektrinus is narrow like in ants. The cuneal fracture in H- elektrinus runs diagonally to the long body axis while in H. succinus it crosses at a right angle. There is also a difference in hair distribution on head (although it can be attributed to the condition of a specimen preserved in amber). Setae in H. elektrinus are much denser on genae, clypeus and first and second antennal segments. 15010Polskie Pismo Entomologiczne 69 (2)

Fig. 3. Hallodapomimus succinus, holotype female, dorsal view.

Fig. 4. Hallodapomimus succinus, female, lateral view. HERCZEK A.: First Phylinae (Heteroptera: Miridae) from Baltic amber 151

The colouration of both species is almost identical, and stable in both sexes of H. elektrinus.

DISCUSSION

Two distinct tribes ofthe subfamily Phylinae: Hallodapini and Leucophoropterini, have no any precise morphological feature (not connected with genitalia) attributable only to one of them. Hence, a number of morphological characters occur in representatives of both tribes, even though quite frequently a characteristic feature of a genus or species of one tribe appears much more rarely in some representatives ofthe other (SCHUH 1974). Among the characters present in species of both tribes there are: frequent similarity to ants, the presence of pale patches on dark hemelytra, flattened collar, frequent narrowing of the anterior part of pronotum, parallel, usually hair-like parempodia, and very small pulvilli. The differences include: usually convex head in the hind part in Hallodapini, which in representatives of Leucophoropterini it is often concave; scutellum is flat in Leucophoropterini while in Hallodapini it is torous or spike-shaped. The species of Leucophoropterini have narrow abdomen. Some features ofthe species described from Baltic amber resemble characters observed in representatives of the two tribes. These are: wide and flattened collar, pronotum narrowed in the anterior part, abdomen narrowed basally in females, the distal margins of hemelytra weakly notched, characteristic creamy patches on dark brown corium, parallel, hair-like parempodia and very small pulvilli. The head rounded in the hind part, the position of eyes close to the posterior margin of pronotum, and torous scutellum suggest that Hallodapomimus should be placed within the tribe Hallodapini. Within Hallodapini six generic groups are distinguished (SCHUH 1974). Among them Cremnocephalus FIEB. has long first tarsal segment, which is uncommon feature among Phylinae (WAGNER 1970). A similar structure is also observed in representatives of the genus Myrmicomimus REuT., which has a different colour pattern on hemelytra. The new genus Hallodapomimus described from Baltic amber, like recent genera Hallodapus and Myrmicomimus, has long first tarsal segment, twice as long as the second one. However, white patches on hemelytra are totally different than in the recent genus Cremnocephalus FIEB. Also a collar is dissimilar, flat in the amber genus and rounded in Cremnocephalus FIEB. and Myrmicomimus RUET. A torous scutellum and the arrangement of pale patches may suggest that the discussed genus belongs to the group Systellonotus FIEB., however the convex hind part of head - concave in the species of Systelonotus - contradicts this suggestion. It is also difficult to place the described genus within the group Hallodapus FIEB. characteristic in having stridulatory structures and eyes in males and females of similar size. In females of Hallodapomimus eyes are smaller than in males, but the difference is not as significant as in recent representatives of Hallodapus FIEB. Characters observed in Hallodapomimus preclude its placement within the remaining groups: 152 Polskie Pismo Entomologiczne 69 (2)

Aoelocoris REuT., Formicopsella Popp. and UHLER (they lack some of features, such as: the absence of pulvilli or pulvilli long, pale patches on hemelytra, dorsoventrally elongated head, eyes removed from the posterior margin ofpronotum etc.). The presence of a long first tarsal segment in the amber genus Hallodapomimus and a pattern on hemelytra different than in the recent genus Cremnocephalus FIEB. may confirm SCHUH'S (1974) suggestion, that this character is not stable within Phylini.

REFERENCES

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Received: 12 April 2000