Cockroaches Probably Cleaned Up after Dinosaurs

Dinosaurs undoubtedly produced huge quantities of excrements. But who cleaned up after them? Dung beetles and flies with rapid development were rare during most of the Mesozoic. Candidates for these duties are extinct cockroaches (Blattulidae), whose temporal range is associated with herbivorous dinosaurs. An opportunity to test this hypothesis arises from coprolites to some extent extruded from an immature cockroach preserved in the amber of Lebanon, studied using synchrotron X-ray microtomography. 1.06% of their volume is filled by particles of wood with smooth edges, in which size distribution directly supports their external pre-digestion. Because fungal pre-processing can be excluded based on the presence of large particles (combined with small total amount of wood) and absence of damages on wood, the likely source of wood are herbivore feces. Smaller particles were broken down biochemically in the cockroach hind gut, which indicates that the recent lignin-decomposing termite and cockroach endosymbionts might have been transferred to the cockroach gut upon feeding on dinosaur feces

Globula Vršanský 2009, n. gen.

Genus <i>Globula</i> n. gen. <p> ETYMOLOGY. — After <i>globus</i> (Latin for globe) – alluding to the large globular head.</p> <p>DIAGNOSIS. — Head extremely large, globular. Apical femoral spur extremely long and sharply curved. Coxa with reticulations (Fig. 7C).</p> <p>DESCRIPTION</p> <p> Head large, globular, with eyes projected beyond the head outline. Antenna very long (1.5 times longer than the total body length), with long <i>sensilla chaetica</i> in several rows. Palps of short-type (characteristic of the Blattulidae), but unusually long, with the third segment curved. Legs thin and slender, the hind leg with very robust and long, curved apical spur. Tarsi and cerci extremely long.</p> <p>REMARKS</p> <p> The present taxon is preliminary categorized within Blattulidae based on characteristic antennal <i>sensilla chaetica</i>, large globular head, short and robust palps. Nevertheless, the curved second segment of palps, slender hind legs and extremely long apical femoral spur are characters unknown in any Blattulidae, and thus the attribution within some other families (Caloblattinidae or some unknown family) cannot be definitively excluded. Moreover the extremely long slender cercus is uncharacteristic of the Blattulidae. The erection of a new family based on unperfectly preserved nymph appears not supported enough.</p> <p> If belonging to the Blattullidae, the present taxon is well established basing on the abovementioned differences, and can be related only to the <i>Nula</i> Vršanský, 2008 from the Cenomanian Sisteron amber based on the reticulations of the coxa. On the other hand, <i>Nula</i> is a standard blattulid with a completely different habitus with head of a standard size. The coxa reticulations may be eventually plesiomorphic and characteristic for the Caloblattinidae (structure of coxae are unknown in Caloblattinidae) or some unknown family.</p>Published as part of <i>Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1)</i> on pages 80-81, DOI: 10.5252/g2009n1a7, <a href="http://zenodo.org/record/5380613">http://zenodo.org/record/5380613</a&gt

Sivis Vršanský 2009, n. gen.

Genus <i>Sivis</i> n. gen. <p> TYPE SPECIES. — <i>Sivis odpo</i> n. sp., by monotypy.</p> <p> ETYMOLOGY. — After <i>si vĪs</i> (Latin for “if you like it”).</p> <p> DIFFERENTIAL DIAGNOSIS. — The present taxon may be categorized within the Mesoblattinidae based on modern morphology of head with globular eyes and clypeus, with strong antenna bearing a row of short <i>sensilla chaetica</i>, by the derived form of pronotum and long marginal chaeta on head, nota and abdomen.</p> <p> The present genus may be differentiated from <i>Praeblattella</i> Vršanský, 2003 based on the characteristic pronotum coloration, which is sophisticated in <i>Praeblattella</i>.</p> <p> <i>Mesoblattina,</i> <i>Archimesoblatta</i> Vršanský, 2003 and <i>Hispanoblatta</i> Martínez-Delclós, 1993 differ by having a simpler form of pronotum (plesiomorphy).</p> <p> The derived form of pronotum is synapomorphic with the <i>Praeblattella</i>, autapomorphic in sophisticated pronotal coloration. The pronotum of <i>Hispanoblatta</i>, <i>Mesoblattina</i> and <i>Archimesoblatta</i> Vršanský, 2003 is plesiomorphic. <i>Brachymesoblatta</i> Vršanský, 2003 belongs to the Liberiblattinae. Thus, <i>Sivis</i> n. gen. may be the sister genus of <i>Praeblattella</i>.</p> <p>DESCRIPTION</p> <p> Head, notum and abdomen with distinct very long marginal <i>sensilla chaetica</i>. Head large, with dark globular eyes protruding beyond the head outline. Antenna very wide, with long scape, shorter pedicel and very long third antennal segment. Mandible very large, with numerous small sharp teeth. Maxillary palp with cup-like apical segment. Pronotum almost quadrate in outline, body not wide.</p> <p>REMARKS</p> <p>Distinct cup-like apical segment of maxillary palp is a phylogenetic novelty, absent in Mesozoic families. The terminal segment may be cup-like in the pollinating Skokidae, but not as distinct as here.</p>Published as part of <i>Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1)</i> on page 86, DOI: 10.5252/g2009n1a7, <a href="http://zenodo.org/record/5380613">http://zenodo.org/record/5380613</a&gt

Sobytie Vršanský, 2010, gen.n.

Sobytie gen.n. Type species. Sobytie tungusicum sp. n., by monotypy. Differential diagnosis. The present taxon differs from all other representatives of the family in having comparatively short Sc; expanded R with differentiated RS reaching apex, main M stem directed anteriorly, well organized V-shaped cross-veins of both wings; hindwing with a polyphagoid scheme of venation (remigium-supporting A 1 with basal short branches). Description. Both wings with distinct, mostly straight but locally zig-zag intercalaries on the lower surface of convex membrane folds (fig. 1 B, 2 D). Distinct cross veins present in both wings, changing into close network locally in clavus and hindwing CuA area. Forewing costal field wide, Sc branched. R 1 and RS weakly separated, but RS present. M expanded, branches straight or very slightly arched. CuA with anteriormost branch further branched. Anal veins branched. Hindwing with simplified venation, according to the presence of curved, branched A 1 with short blind branches in the remigium, the pleating was likely not fan-like. Sc very strong but short. R 1 very short, branched terminally, RS branches also short. Media reduced to few veins. CuA secondarily branched, with stochastical occurrence of fenestrate areas between intercalaries and cross-veins. A 1 with short basal branches and long, nearly straight anteriormost branch. Remarks. There are small (forewing length 9–13mm), similar representatives of the genus Phyloblatta, known from the Permian of Elmo in U.S.A. and Obora in Czech Republic; P. compacta (Sellards, 1908); Schneider 1984, pl VIII, figs. 9–10. Nevertheless, the shorter Sc, expanded R, different organization of M and CuA branches (see diagnosis) and much better organized V-shaped cross-veins are good arguments to distinguish the new genus. The hindwing with remigium-attached A 1 differs from the Phyloblatta, which does not have basal blind branches. Otherwise the hindwing can be compared with some Permocarboniferous Phyloblattidae (Schneider 1984) (for phylogenetic relations see discussion). RS is apparently (fig. 2 b) part of R and not M merged to the distal part of R (as in Schneider 1977, 1983), because of a regular venation in that area – this pattern is characteristic for phyloblattid descendants (unless RS is actually a merged cockroach M in general). Zig-zag intercalaries forming a secondary corrugation (see Schneider 1984 a, pp. 9–14, pls. 2, 5) appear increasingly from the Early Permian and are typical for most Late Permian phyloblattids. Intercalaries are distributed on the bottom side of the membrane, which rises over the wing profile near them (see fig. 1 B). Etymology. The genus name is the transliteration of обытие (Russian for event); gender neuter.Published as part of Vršanský, Peter, 2010, A new genus and species of cockroach (Blattida: Phyloblattidae) from the Permian / Triassic boundary beds of Tunguska Basin in eastern Siberia, Russia, pp. 55-61 in Zootaxa 2353 on pages 56-57, DOI: 10.5281/zenodo.27564

Mesoblattinidae Handlirsch 1906

Family MESOBLATTINIDAE Handlirsch, 1906 DIAGNOSIS. — Head globular, eyes projected more above the outline of the head. Antennal segments with few very short subapical sensilla chaetica. Maxillary palp of mediate length. Body plain, cerci with about 15 segments without long sensilla chaetica; styli absent. Female with internalized, sometimes reduced external ovipositor. Wings with round dark veins and pale round intercalaries. Sc simple or with max. 3 branches; RS differentiated; A branched. Hindwing with sophisticated, comb-like R1, differentiated RS; simple M; branched CuA, simple CuP and simple A1. Pleating veer-like.Published as part of Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1) on page 86, DOI: 10.5252/g2009n1a7, http://zenodo.org/record/538061

Liberiblattinidae Vrsansky, MOTS CLES 2002

Family LIBERIBLATTINIDAE Vršanský, 2002 DIAGNOSIS. — Head globular, eyes projected more or less within outline of the head. Antennal segments bearing median short sensilla chaetica. Maxillary palp robust.Body thick, cerci with few segments. Female with internalised or short external ovipositor.Wings with round dark veins and more pale round intercalaries, usually strongly curved (also locally, with local irregularities), with cross-veins. Sc very long, branched; RS not differentiated; A simple or branched. Hindwing with simply branched R1, differentiated RS, with secondarily branched CuA, simple CuP and curved and branched A1. Pleating not veer-like.Published as part of Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1) on page 83, DOI: 10.5252/g2009n1a7, http://zenodo.org/record/538061

Eadiidae Vršanský 2009, n. fam.

Family EADIIDAE n. fam. TYPE GENUS. — Eadia n. gen. OTHER GENUS INCLUDED. —? Raphidiomimula Grimaldi & Ross, 2004 from the Upper Albian Burmese amber. RANGE. — Eurasia, Upper Albian. DIFFERENTIAL DIAGNOSIS. — The family may be included within Dictyoptera based on the general habitus, hypognathous head, multisegmented, partially fusiform cerci and wing pads. It can be further categorized as belonging to cockroaches and not to mantises based on the generally short body, short pronotum and particularly cursorial forelegs with short tibiae (but with two rows of spurs like in mantises and liberiblattinid cockroaches), and extremely long tarsi. Eadiidae n. fam. differs from all known cockroach families in having hypognathous, but largely expanded and prolonged head, extremely short pronotum, by curved carination and long tarsi of forelegs and by extremely elongated cerci. DESCRIPTION Head hypognathous, largely prolonged. Eyes (with small facets) protruding beyond the head outline. Three ocelli present. Pronotum extremely short and wide. Wing pads distinct. Legs robust, forelegs tibiae short, with sharply curved spurs, fore tarsi extremely long. Cercus extremely long, and in the apical half extremely narrow. REMARKS The extremely long extremities, and long narrow maxillary palps resembling those of families Caloblattinidae and Raphidiomimidae (symplesiomorphies), exclude affinities with Polyphagoidea (including Liberiblattinidae and Skokidae), Umenocoleoidea and Blattuloidea. The generally short body and elongated fore tarsi are strong autapomorphies, the former eventually serving for capturing prey. Another possible synapomorphy with primitive mantises is the elongated cercus. The composition of head is a strong autapomorphy, a trend towards prolongation perhaps inherited from the Caloblattinidae, whose heads are diverse. Raphidiomimidae have prognathous head (autapomorphy)and very short fore tarsi(plesiomorphy).Thus, the new family may be attributed within Caloblattinoidea and derived directly from the Caloblattinidae. Even when the morphology of head appears identical with the Raphidiomimula, the categorization of this genus is obscure, because of its longer pronotum and shorter tarsi.Published as part of Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1) on page 82, DOI: 10.5252/g2009n1a7, http://zenodo.org/record/538061

Sobytie tungusicum

Sobytie tungusicum gen. et sp.n. (Figs. 1–2) Holotype. PIN 2010 / 16 (14). Part and counterpart of all four wings (two complete). Russia, Krasnoyarsk Territory, Bugarikta River, Bugarikta Formation (lower part). Description. Forewing length 12 mm. Sc with 6 veins meeting margin, one Sc branch dichotomised. R 1 with 11, RS with 7 veins ending at margin. The basalmost R branched terminally. M and CuA each ending with 7 veins at margin. Diagonal kink (pseudovein in clavus) separating the posteriormost third of clavus distinctly. Characteristic V-shaped reticulation apparent in cubital, medial and the two most apical radial areas. Main veins dark, appearing black when compared with paler brown intercalaries and rest of wing. Coloration monochromatic, probably pale brown. Hindwing Sc simple. R 1 is only indicated by the basalmost branch with 4 veins at margin; RS expanded, with 9 veins meeting margin. M reduced to 4 veins. CuA with seven veins at margin and an additional blind branch. A 1 with 3 basal branches. Hindwing without coloration. Derivation of name. The specific name tungusicum is derived from the major river of the region, the Nižnaja Tunguska River.Published as part of Vršanský, Peter, 2010, A new genus and species of cockroach (Blattida: Phyloblattidae) from the Permian / Triassic boundary beds of Tunguska Basin in eastern Siberia, Russia, pp. 55-61 in Zootaxa 2353 on pages 57-59, DOI: 10.5281/zenodo.27564

Eadia Vršanský 2009, n. gen.

Genus Eadia n. gen. TYPE SPECIES. — Eadia aidae n. sp., by monotypy. DIFFERENTIAL DIAGNOSIS. — Head like in Raphidiomimula Grimaldi & Ross, 2004 (whith temporal space invisible, and thus the presence of ocelli not validated). Whole body considerably shorter, and robust, cerci more derived, elongated with narrow, almost filiform distal half. Fore tarsi considerably longer. ETYMOLOGY. — After eādem, Latin for “by the same way”. DESCRIPTION Eyes large, protruding beyond the head outline. Facets very small. Space between eyes very wide. Three ocelli are plain and pale, covered by cuticle. Three rows, each with 10 short chaeta, present above clypeal sutura. Maxillary palp long and narrow, with sparse slender sensilla chaetica. Fore femur with two distinct rows of spurs, tibia with very strong, curved spurs. Tarsi very long (longer than tibiae), with numerous small sensilla chaetica. Cerci long, of standard general fusiform habitus in basal half, but extremely thin and elongated apically. REMARKS The robust and considerably shorter body, and even more elongated cerci are strong autapomorphies.Published as part of Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1) on pages 82-83, DOI: 10.5252/g2009n1a7, http://zenodo.org/record/538061

Blattulidae Vishniakova 1982

Family BLATTULIDAE Vishniakova, 1982 DIAGNOSIS. — Usually small species (forewing usually well under 10 mm, extremely rarely 17 mm). Head globular, eyes projected more or less within outline of the head. Occiput usually with two distinct dark stripes. Antennal segments bearing a set of transversal ridges and five to seven subapical sensilla chaetica about as long as half segment length. Maxillary palp short and robust. Body thick, cerci with few segments bearing extremely long sensilla chaetica; styli present. Female with short external ovipositor. Wings with plain dark veins and more pale plain intercalaries, usually with cross-veins. Sc simple; RS not differentiated; A simple. Hindwing with comb-like R1, differentiated RS, usually with simply branched CuA, simple CuP and curved and branched A1. Pleating is not veer-like.Published as part of Vršanský, Peter, 2009, Albian cockroaches (Insecta, Blattida) from French amber of Archingeay, pp. 73-98 in Geodiversitas 31 (1) on page 75, DOI: 10.5252/g2009n1a7, http://zenodo.org/record/538061