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To Possible Assignment of a New Genus of Lamellicorn Beetles from the Lower Cretaceous of Mongolia to the Subfamily Pleocominae (Coleoptera, Scarabaeidae) G

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To Possible Assignment of a New Genus of Lamellicorn Beetles from the Lower Cretaceous of Mongolia to the Subfamily Pleocominae (Coleoptera, Scarabaeidae) G Paleontological Journal, Vol. 36, No. 3, 2002, pp. 279–282. Translated from Paleontologicheskii Zhurnal, No. 3, 2002, pp. 51–55. Original Russian Text Copyright © 2002 by Nikolajev. English Translation Copyright © 2002 by åÄIä “Nauka /Interperiodica” (Russia). To Possible Assignment of a New Genus of Lamellicorn Beetles from the Lower Cretaceous of Mongolia to the Subfamily Pleocominae (Coleoptera, Scarabaeidae) G. V. Nikolajev Al-Farabi State University of Kazakhstan, ul. Vinogradova 95, Almaty, 480012 Kazakhstan Received March 31, 2000 Abstract—A new genus, Crecotoma gen. nov., representing a separate tribe in the subfamily Pleocominae, is described from the Lower Cretaceous of Mongolia. The systematic position of this subfamily and phylogenetic links with other taxa of Scarabaeidae are discussed. INTRODUCTION genera Acoma and Benedictia showed that they should be removed from the Pleocominae (Ritcher, 1969; The venation of the costal area of the wing is clearly Hardy, 1978); thus, this subfamily is monotypic in the traced in a fairly complete fossil beetle from the Lower modern fauna. Cretaceous of Mongolia, which is kept in the collection of the Paleontological Institute, Russian Academy of In the entire superfamily of the lamellicorn beetles, Sciences (PIN) (Figs. 1a, 1b). Distinctive features of the structure of the fore coxal pits, which are unclosed, the venation allow easy identification of this beetle it is unique to the genus Pleocoma. In my opinion, the easily as a species of the superfamily of the lamellicorn taxonomical rank of the group and its place in the sys- beetles. The compression represents a female, since the tem of the superfamily should be based on this feature. traces of the external female genitalia; namely, the cox- If the structure of the fore coxal pits of Pleocoma is ites with well-preserved and fairly large styles are a plesiotypic feature, this taxon should be considered to clearly visible. Six visible sternites are developed in the be a sister group with respect to all other suprageneric abdomen of the beetle. The fore tibia has three outer taxa of the superfamily taken together. This conclusion teeth, and the middle and hind tibiae bear one trans- inevitably comes from the analysis of other morpholog- verse ridge each. The reduction of the vein R3 in the ical peculiarities of the imago. This genus retains only apical part of the wing occurs among the Recent lamel- a few plesiomorphies (mainly in the structure of the licorn beetles only in the monotypic subfamily Pleo- antennae), but acquires a number of apomorphies (see cominae, a small relict group, which is now endemic to below). These apomorphies are to be treated as autapo- North America. Species of this genus have this vein in morphies, and the rank of the taxon will be in no case a form of a stripe, which is sclerotized somewhat stron- less than familial. ger than the rest of the wing (Fig. 1d). By assuming that the open fore coxal pits of Pleo- However, the structure of the external female geni- coma is an apotypic feature, one may try to find out talia with coxites bearing large styles is not characteris- which of the generic apomorphies may represent syna- tic of Recent Pleocominal species (Zunino, 1984, text- pomorphies with other taxa of the superfamily, since fig. 4). This important feature makes us consider this the majority of other characters of these beetles are not new beetle to be a representative of a separate taxon of uncommon among other suprageneric taxa of lamelli- a suprageneric rank. The habitus resembling that of corns. In this case, the set of features characteristic of phytophagous lamellicorns from the Pleurosticti spe- this genus will only suffice to consider it to be a sister- cies group and the presence of some apomorphies com- group of one or several taxa rather than of all lamelli- mon to the Pleocominae and the species under study corn beetles together; the suprageneric rank of the (the absence of a leathery bordering at the apex of the taxon should be at least one level lower than the rank pronotum, one transverse ridge on the middle and hind adopted for all lamellicorn beetles. This is either a tribe tibiae, and adjoining spurs on the same tibiae) allow us (subfamily) within the family or a family within the to treat them as synapomorphies and to rank this taxon superfamily. as a tribe in the subfamily Pleocominae. The genus Pleocoma is characterized by the follow- The Recent genera Pleocoma LeConte, 1856, ing features (the plesiomorphic state is designated by p, Acoma Casey, 1881, and Benedictia Sanderson, 1939 the apomorphic state, by a). were included in the subfamily Pleocominae (Arnett, The antennae are 11-segmented (p) with a multiseg- 1963). However, the investigation of characters of the mented club (p); the mouthparts (labrum and mandi- 279 280 NIKOLAJEV (a) (b) (c) (d) (e) Fig. 1. Fossil and Recent Scarabaeidae: (a)–(c) Cretocoma tologoica sp. nov.: (a) and (b) holotype PIN, no. 4271/217: (a) top view, (b) bottom view, (c) reconstruction; (d) and (e) wing venation in Recent species: (d) Pleocoma australis Fall., (e) Oncerus sp. The arrows indicate (b) the remains of the external female genitalia and (d) and (e) the position of the veins R3. Scale bar 1 mm. bles) are hidden under the clypeus (a); the leathery bor- To illustrate this, it is suffice to consider the structure dering along the anterior margin of the pronotum is of antennae and wing venation. Retaining the vein R3 absent (a); the wings are developed only in males (a); (Fig. 1e) in the apical portion of the wing (plesiomor- the vein R3 is reduced (a); two free veins are present phy), the subfamilies Pachypodinae and Oncerinae pos- between the cubital vein and the first anal vein con- sess antennae with a smaller number of segments than nected with the wing base (p); the abdomen has six vis- those belonging to the Pleocominae (apomorphy). ible sternites (p); abdominal spiracles are situated on A multisegmented antennal club of the Pachypodinae, the intersegmental membrane between the tergites and which in turn is a plesiomorphy against a three-seg- sternites (p); the fore tibia has more than three teeth on mented club of the Oncerinae, goes with a smaller num- its outer margin (p); the middle coxae are not widely ber of antennomeres (autapomorphy) compared with separated (p); the middle and hind tibiae have one the ten-segmented antennae of the Oncerinae. transverse ridge on their outer face (a) and contiguous At present, I do not think it sensible to consider this apical spurs (a); and the coxites of the external female phyletic line to be a single taxon, i.e., a subfamily genitalia bear very small styles (a). within the family Scarabaeidae or a family within the A number of the above apotypic features of Pleo- superfamily Scarabaeoidea, and, thus, to lower the rank coma, i.e., the labrum and mandibles hidden under the of each group constituting the cladon; it should be clypeus and the middle and hind tibiae having one noted that the name Pachypodinae Erichson, 1840 has transverse ridge on their outer side, are characteristic of a priority. the oligotypic subfamily Oncerinae Saylor, 1938 and A founder species of this line must have the whole monotypic subfamily Pachypodinae Erichson, 1840 set of characters of the genus Pleocoma except for the [authors of the taxa and years of publication are given open anterior coxal pits, contiguous spurs on the hind after Ritcher (1969) and Laurence and Newton (1995)]. tibiae, wing venation in the apical portion with the vein Currently, I see no insurmountable obstacles to con- R3 that must be developed, small styles on the coxites sider these taxa to be members of one phyletic line. of the external female genitalia, and, most probably, However, a combination of plesiotypic and apotypic wing reduction in female. Wing reduction only in the characters in each of these taxa (Pachypodinae, Pleo- female may be caused by unfavorable conditions of the cominae, and Oncerinae) is such that each subfamily life of larvae that lead to high mortality rates during this may be a sister-group of any member of this phyletic line. stage (long larval periods or developing in arid and PALEONTOLOGICAL JOURNAL Vol. 36 No. 3 2002 TO POSSIBLE ASSIGNMENT OF A NEW GENUS OF LAMELLICORN BEETLES 281 semiarid conditions, Nikolajev, 1984). Such a reduction (also apomorphy), the presence of well developed wings is called ootrophic and most probably developed quite in female (plesiomorphy), and in the presence of well recently in two of the three above-mentioned groups, developed large styles of the gonocoxites of female i.e., subfamilies Pachypodinae and Pleocominae; unfor- genitalia (plesiomorphy). Regrettably, some details, tunately, I do not know whether the ootrophic reduction such as the position of the labrum and the mandibles of wings is developed in Oncerinae females. against the clypeus (usually, this feature is well pre- Possible relations of Pleocominae with species of served in fossils) and whether or not the anterior coxal the families Diphyllostomatidae, Lucanidae, and Pas- pits are closed (this feature is hardly visible in lamelli- salidae are thoroughly studied in (Nikolajev, 2000). corn beetle fossils), cannot be observed. This taxon Possible assignment of the fossil genus Proteroscar- may be a sister group of the Pleocomini sensu stricto abaeus Grabau, 1923 from the Mesozoic of China to despite the state of these characters in representatives of the subfamily Pleocominae has also been hypothesized the Cretocomini. (Nikolajev, 1996). A multisegmented antennal club and fore tibiae with a large number of teeth along the outer Genus Cretocoma Nikolajev, gen. nov. margin make it possible to bring this genus together with the type genus of the subfamily.
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