N. Jb. Geol. Paläont. Abh. 2008, vol. 247/3, p. 353–381, Stuttgart, March 2008, published online 2008

The enigmatic Mesozoic insect taxon Chresmodidae (Polyneoptera): New palaeobiological and phylogenetic data, with the description of a new species from the Lower Cretaceous of Brazil

Xavier Delclòs, Barcelona, André Nel, Paris, Dany Azar, Fanar-Matn, Günter Bechly, Stuttgart, Jason A. Dunlop, Berlin, Michael S. Engel, Lawrence, and Sam W. Heads, Portsmouth With 10 figures

DELCLÒS, X., NEL, A., AZAR, D., BECHLY, G., DUNLOP, J. A., ENGEL,M.S.&HEADS, S. W. (2008): The enigmatic Mesozoic insect taxon Chresmodidae (Polyneoptera): New palaeobiological and phylogenetic data, with the description of a new species from the Lower Cretaceous of Brazil. – N. Jb. Geol. Paläont. Abh., 247: 353–381; Stuttgart.

Abstract: The morphology of the enigmatic, Mesozoic, aquatic insect family Chresmodidae is redescribed and its phylogenetic affinities among the polyneopterous orders discussed. Study of the complete venation of both fore- and hind wings observed in some specimens from the Spanish Barremian, permit us to postulate the hypothesis that the family belongs to the Archaeorthoptera, thus to the orthopteroid lineage rather than to crown-group Phasmatodea or to the more inclusive group Holophasmatodea (sensu GRIMALDI &ENGEL, 2005). New specimens from Spain, Lebanon, Brazil, and Germany permit a new re-description of some chresmodid body structures with concomitant implications for the phylogenetic position of the family. Chresmoda neotropica n. sp. is described from the Aptian-Albian of the Crato Formation (northeast Brazil). The functional morphology eschweizerbartxxx proposed for some of their specialized structures suggest a new hypothesis of Chresmoda palaeo- biology, and related to this some implications for the localized palaeoenvironment as well as global palaeoclimate. The problematic Sternarthron spp. from the Upper Jurassic of Solnhofen were described as probable palpigrades (Arachnida: Palpigradi), based on type material originally thought to be fossil insects. The affinities of Sternarthron HAASE, 1890 have been questioned. Our restudy of HAASE’s types clearly confirmed earlier assumptions that these fossils represent nymphal specimens of chresmodids. Consequently, Sternarthron has to be considered as an invalid junior synonym of the fossil insect genera Propygolampis WEYENBERGH, 1874 and Chresmoda GERMAR, 1839.

Key words: Insecta, Polyneoptera, Archaeorthoptera, Sternarthronidae, Chresmodidae, Palpigradi, Functional Morphology, Phylogeny, Paleoecology, Mesozoic, Crato Formation, Solnhofen Litho- graphic Limestones.

1. Introduction and historical context tarsi with more than 40 tarsomeres (Fig.7.2), a feature which is unique within Insecta (NEL et al. 2004). The Chresmodids are large fossil insects with a water- forelegs are usually directed anteriorly, while the strider-like habitus. They have short and thick middle and hind legs are directed latero – posteriorly. antennae and prognathous chewing mouthparts with They have two pairs of membranous wings with a strong mandibles, and large compound eyes. The legs slender forewing with long and parallel longitudinal are extremely prolonged with very long femora, veins and a broad anal fan in the hind wings, typical of shorter tibiae, and long, multi-segmented, flagellate polyneopterous insects (Figs.6.3, 8). There are long,

DOI: 10.1127/0077-7749/2008/0247-0353 0077-7749/08/0247-0353 $ 7.25 © 2008 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart 354 X. Delclòs et al. monomerous, lancelolate cerci at the end of the ab- He considered the types of both P. bronni and C. domen (similar to Phasmatodea), and females have a obscura to be unusable, since the original fossils had prominent orthopteroid ovipositor. Males seem to be been painted in (a ‘bemalter Artefact’), but gave no apterous. The nymphs are similar to the adults except justification for rejecting Propygolampis as a valid for a much smaller size, a distinctly shorter abdomen, name and substituting his own name, Halometra;now and the absence of wings. Consequently the deve- regarded as a junior synonym of Propygolampis (e.g. lopment of chresmodids was heterometabolous as CARPENTER 1992). OPPENHEIM, like GERMAR, thought in orthopterans and other polyneopterous insects. H. gigantea was a semiaquatic hemipteran, referring Chresmodids probably lived on the surface of lakes the genus to the family Hydrometridae. In addition and lagoons (contra BAUDOIN 1980 who did not know to this, he described a further Solnhofen species, the true structure of chresmodid legs), and fed on Halometra ? minor OPPENHEIM (1888) based on three insects and other small animals at the water surface, smaller fossils in the Bayerische Staatssammlung für just like the unrelated, but superficially similar, Paläontologie und Geologie, Munich. Although modern Gerridae. Because of their significant larger OPPENHEIM (1888) was uncertain about the generic size and weight it cannot be excluded that chresmo- affinities of H.? minor, he specifically noted that the dids might have needed floating water plants near overall shape and number of legs (i. e., six) meant the shore to support their heavy bodies on the water that they could not be crustacean larvae or arachnids. surface. The group is exclusively Mesozoic and may CARPENTER (1992) propagated the younger synonym have suffered extinction during the Late Cretaceous Propypolampis WEYENBERGH, 1874 as the valid name greenhouse climate. for the fossils previously known as Chresmoda Since GERMAR (1839) named Chresmoda obscura GERMAR, 1839, and mentions a paper by himself for an insect found in the Upper Jurassic of Soln- (listed as “in press” for 1992) that allegedly revealed hofen-Eichstätt, the taxonomic placement of this the holotype of Chresmoda to be a locust and was only enigmatic species has been dramatically reinterpreted later confused with the “fossil water striders”. There- several times. GERMAR (1839) originally established fore, CARPENTER (1992: 181) classified Chresmoda the species for a “flattened-like shaped insect, with a within Orthoptera-Caelifera as a “little-known genus pronotum and with long hind legs”, and placed it probably related to Acrididae”, while he considered among the mantises (Mantodea). In the same paper he Propygolampis to be related to Phasmatodea. How- established also the species Pygolampis giganteaeschweizerbartxxx , ever, the cited paper of CARPENTER that was sup- which was a larger insect, similar to C. obscura,but posedly “in press” in 1992 in the journal Psyche was with forelegs shorter than the mid- and hind legs. never published (FURTH 1994) and the manuscript has GERMAR believed P. gigantea was a hemipteran to be considered as lost because it is neither archived similar to the living water-strider Gerris. This paper with the intended publisher nor could it be found introduced an immediate nomenclatural problem since among CARPENTER’s archives in his laboratory at the GERMAR actually ascribed both species names to his MCZ (GB and MSE, pers. obs.). compatriot, MÜNSTER, although MÜNSTER was not an HANDLIRSCH (1906-1908) proposed that Pygo- author on this paper and the work is clearly that of lampis gigantea was a junior synonym of Chresmoda GERMAR. Subsequent authors (e.g. CARPENTER 1992) obscura, and established the new family Chresmo- have correctly treated GERMAR as the author of these didae and providing a discussion about its inclusion taxa. WEYENBERGH (1869: 27) listed “Pygolampis in Phasmatodea or Orthoptera. Subsequently, HAND- gigantea GERM.” and mentioned three beautiful fossils LIRSCH (1925, 1926-1930) considered chresmodids as of this species from the Teyler Museum in Haarlem, of aquatic phasmatodeans. Unfortunately, the holotype which he figured one (pl. 2, fig. 21) with the catalogue of C. obscura is missing, whereas the holotype of P. number 6395. Subsequently, a similar fossil hemi- gigantea is housed in the Munich Museum (Fig. 2.1). pteran genus Propygolampis WEYENBERGH, 1874 was HANDLIRSCH (1906-1908) placed Gryllidium oweni created for a new Solnhofen species, Propygolampis WESTWOOD, 1854 into Chresmoda. The latter is an bronni WEYENBERGH, 1874. Meanwhile, ASSMANN isolated forewing with longitudinal veins and with (1877) synonymised P. gigantea with C. obscura, re- dense crossveins and basal archaedyction, the holo- garding them as orthopterans. OPPENHEIM (1888) later type of which is now lost (SWH pers. obs.). synonymised P. gigantea, C. obscura, and P. bronni MARTYNOV (1928) placed the suborder Chresmo- under the name Halometra gigantea (GERMAR, 1839). dodea into the Phasmatodea, including four families The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 355 related by their wing venation: Chresmodidae (Trias- Orthopterida (i.e., secondarily autapomorphic for sic-Cretaceous), Aeroplanidae (Triassic), Necrophas- Phasmatodea). Some authors place them close to the matidae, and Aerophasmidae (both Jurassic). ESAKI Embiodea, or even with modern Grylloblattodea and (1949) established the species Chresmoda orientalis Mantophasmatodea (see discussion below). More for an insect with short body (24 mm), but with long critical work on the phylogenetic relationships of legs and antennae (11 mm), from a specimen found Phasmatodea and putative relatives is required. in the Upper Jurassic of Ta-hsing-fang-tzu, Jehol in Numerous discoveries in the Upper Jurassic of China (this specimen is also missing today). Germany and Lower Cretaceous of Mongolia, China, SHAROV (1968) considered that chresmodids were Brazil, Spain, and Upper Cretaceous of Lebanon, phasmatodeans and elevated the group to superfamily provide new evidence for the phylogenetic placement rank, as Chresmodidaea, including therein the Meso- of the Mesozoic family Chresmodidae, questioning zoic Chresmodidae and Recent Phasmatidae and some of the traditional views. Phyllidae. POPOV (1980) considered chresmodids as aquatic gerromorphs (Hemiptera: Heteroptera) and 2. The enigmatic genus Sternarthron PONOMARENKO (1985) as an unusual lineage of para- HAASE, 1890 plecopterans. MARTÍNEZ-DELCLÒS (1991) supported the attribution to ‘Paraplecoptera’ in the absence of Some specimens of Chresmoda from Solnhofen were information from the new material presented herein. previously placed in other groups of arthropods. A Contrary to MARTYNOVA (1962, 1991) we here restudy of some of them demonstrate close relation- restrict the Chresmododea to the family Chresmo- ships with different ontogenetic stages of Chresmoda. didae, thus excluding the extinct families Aerophas- This is most notably the case for the genus Sternar- matidae, Necrophasmatidae and Aeroplanidae. It thron (Fig. 1.1.-1.3). cannot yet be totally excluded that some of these taxa Palpigrades are a rare order of small arachnids might be more closely related to Chresmododea than (CONDÉ, 1996 for a recent review), which resemble to Orthoptera or Phasmatodea (see below), but whip scorpions and which are usually thought of as currently there are hardly convincing arguments for rather primitive arachnids. Palpigrades are weakly such a placement. However, WILLMANN (2003), sclerotized and the only reliable fossils are from the followedbyGRIMALDI &ENGEL (2005), polarized Cenozoic (ROWLAND &SISSOM 1980). There is, how- several wing venation characters and proposedeschweizerbartxxx a ever, a much older genus,Sternarthron HAASE (1890b) cladogram of putative stem-group members belonging from the Upper Jurassic of Germany which was to the extant Phasmatodea, which included Aerophas- described as an arachnid; most probably a palpigrade. matidae. This supports our restriction of the Chresmo- Despite being created from fossils described as dodea to include only the family Chresmodidae. insects (OPPENHEIM 1888), and soon being placed The phylogenetic relationship of chresmodids was back among the insects (HANDLIRSCH 1906), HAASE’s long disputed and they have been attributed to Hemi- palpigrade interpretation for Sternarthron was ac- ptera (Heteroptera: Gerromorpha), Paraplecoptera cepted by a number of authors (see below), but was (‘Grylloblattodea’ in some systems), Mantodea, not accepted by HARVEY (2003) in his catalogue of the Orthoptera, Polyneoptera of uncertain affinity, and smaller arachnid orders. With a body length of around most often to Phasmatodea. Most recently they 17 mm, Sternarthron is much larger than any living have been considered “unplaced within Gryllones” palpigrade and HAASE’s drawings of it appear some- (RASNITSYN &QUICKE 2002) or within the Ortho- what stylised. Sternarthron was recently placed back pterida (GRIMALDI &ENGEL 2005). NEL et al. (2005) among the insects (CARPENTER 1992) (see below), placed chresmodids among the monophyletic clade although it is not clear that anyone since HANDLIRSCH Archaeorthoptera BÉTHOUX &NEL, 2002, which do (1906) actually restudied HAASE’s type specimens, not correspond exactly to the Orthopterida sensu one of which was clear depicted with four pairs of legs GRIMALDI &ENGEL, 2005, as these authors included by HAASE (1890b: pl. 31, figs 5-6). Meanwhile, other the modern Phasmatodea in their Orthopterida. The Sternarthron fossils have been compared to harvest- Recent Phasmatodea show no visible evidence of men (Opiliones) and sea spiders (Pycnogonida or the pattern of cubito-median veins typical of the Pantopoda) (FRICKHINGER 1999: 37, fig. 54). To Archaeorthoptera, although this could be interpreted address this confusion, we have reviewed the nomen- as an independent, secondary derivation from within clatural history of this material and present a restudy 356 X. Delclòs et al. of HAASE’s types of this very problematic fossil homeomorphic legs, coxal position, lack of a sucking taxon. apparatus and shape of the thorax. HANDLIRSCH (op. HAASE (1890a) disagreed with the previous attri- cit.) accepted that these fossils belonged to the broad butions and considered H.? minor as an arachnid. Sub- group of insects that were at that time included under sequently, HAASE (1890b) redescribed and figured ‘orthopteroids’ and after discussing why they could both Oppenheim’s Munich material (one of which he not belong to groups such as grasshoppers, roaches, figured with eight legs) plus another Solnhofen fossil and mantises, he concluded that they were most simi- housed in the Dresden Mineralogical Museum. He lar to the phasmatodeans (currently placed in their named these Sternarthron, a new genus of fossil own order Phasmatodea). HANDLIRSCH (op. cit.) noted arachnid, recognising two taxa: S. zittelii HAASE, phasmid features in these fossils such as antennal 1890b and S. zittelii var. minus OPPENHEIM, 1888, shape, a thin body, homeomorphic limbs and short, basing this variation on the specimen originally undivided cerci. Although interpreting them as rela- figured by OPPENHEIM (1888: pl. 31, fig.4) which tives of phasmatodeans, and not phasmatodeans in the HAASE regarded as having a broader, less pointed strict sense themselves, HANDLIRSCH still thought that opisthosoma. The name minor became minus to cor- these fossil insects lived on the surface of the water, respond with the gender change in the generic name. (erroneously) noting that some extant phasmatodeans HAASE (1890b) compared Sternarthron to various live underwater. HANDLIRSCH (op. cit.) also studied extant arachnids. HAASE’s spelling zittelii has to be HAASE’s Sternarthron specimens in Munich that he considered as an incorrect original spelling and is here referred to as ‘HAASE’s Spinne’ (which implies a formally emended to zitteli, even though it was mostly spider, but which can, in a broader sense, translate as already used with this correct spelling HAASE sug- arachnid). He noted that it had six legs and the charac- gested that Sternarthron should not be raised to a new teristic cerci and antennae of Chresmoda. On these order, because there was so little material known, and grounds HANDLIRSCH (op. cit.) interpreted the rela- based primarily on his interpretation of a ventral tively small Sternarthron as a nymphal form of what prosoma with a series of divided sternites and an he believed to be a fossil relative of stick insects. opisthosoma with a terminal flagellum, he felt that it HANDLIRSCH’s study and his synonymy of Sternar- was better to place Sternarthron in THORELL’s (1888) thron with Chresmoda appears to have been widely then recently recognized arachnid order Palpigradi. overlooked, at least among arachnologists and re- Finally, although HAASE (1890b: 653) entitledeschweizerbartxxx the last viewers of the Solnhofen fauna. ROEWER (1934) did section of his paper ‘Eine neue Arthrogastren- not accept Sternarthron as a palpigrade, noting the Familie...’, and concludes (p. 657) with a remark differences in size and the number of prosomal about the sternal and palpal morphology charac- sternites and opisthosomal segments, compared to terising the family, at no point in the text, footnotes living palpigrades. He regarded Sternarthron as an or figure legends did he actually propose a family isolated form among the arachnids and discussed evo- name! lutionary aspects of its morphology. Meanwhile, HANDLIRSCH (1906: 525) placed Sternarthron back Sternarthron was occasionally mentioned as a fossil among the insects, specifically placing Propygolampis palpigrade in zoological text-books of the period (e.g. bronni, Halometra gigantea, Halometra? minor, and KÄSTNER 1941; MILLOT 1949). In his fossil arach- Sternarthron zitteli as junior synonyms of GERMAR’s nid monographs PETRUNKEVITCH (1949, 1953, 1955) Chresmoda obscura.HANDLIRSCH (1906) therefore mentioned Sternarthron and accepted it as a palpi- considered GERMAR’s ‘mantid’ and ‘bug’ to be con- grade. It appears that around this time the family name specific, though he figured (HANDLIRSCH 1906: pl. Sternarthronidae was introduced. The oldest record 44, fig. 17) a ‘Chresmoda’ with long, slender legs, we could find was PETRUNKEVITCH (1949: 261) who which resembles GERMAR’s figure of the original P. gave neither an author nor a diagnosis for it. Sub- gigantea far more than it resembles GERMAR’s figure sequently, PETRUNKEVITCH (1955: 115) incorrectly of the type of C. obscura. Furthermore, HANDLIRSCH attributed Sternarthronidae to HAASE (1890) and rejected the interpretation of these fossils (a combi- diagnosed this family on: ‘Thoracic sternites 2 and nation of Chresmoda and Propygolampis material) as 3 separated from each other by intersegmental hemipterans (and specifically Hydrometridae) on the membrane’. grounds of the presence of well developed cerci, CROWSON et al. (1967) included Sternarthron as the multi-segmented antennae, pattern of wing venation, oldest known palpigrade. In reviews of the Solnhofen The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 357

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Fig. 1. 1: 413.1870.VII.45; Sternarthron zitteli (holotype), Upper Jurassic (Tithonian) of Eichstätt (Germany); Bayerische Staatssammlung für Paläontologie und Geologie Museum (BSPGM), Munich. 2: 1964.XXIII.164; Sternarthron zitteli, Upper Jurassic (Tithonian) of Schernfeld (Germany). 3: AS.I.822; Sternarthron zitteli, Upper Jurassic (Tithonian) of Eichstätt. fauna, both KUHN (1961, 1977) and BARTHEL (1978) legs, inferred that it lived in shallow water (a non- accepted Sternarthron as an arachnid. SAVORY (1971) sensical argument given that many fully terrestrial also accepted Sternarthron as a palpigrade, repro- arachnids also have slender legs). ROWLAND &SISSOM duced one of HAASE’s figures and based on the slender (1980: 79) included Sternarthron in a synopsis of the 358 X. Delclòs et al.

Palpigrada and followed PETRUNKEVITCH’s diagnosis a new arachnid order. Consequently, a restudy of of Sternarthronidae. However, they suggested that the types of Sternarthron is long overdue and here overall Sternarthron looked more like a water strider provided. (Hemiptera, Gerridae) than a living palpigrade and Remarks. – The genera Sternarthron, Propygo- that a re-examination of the original material was lampis, Pygolampis, and the species C. obscura comes required to confirm its identity. BARTHEL et al. (1990), from the Upper Jurassic (Tithonian) of Solnhofen, in a revised translation of his original review, regarded Southern Franconian Alb, in Germany. The geological Sternarthron as a rather dubious palpigrade. setting of which was reviewed by BARTHEL et al. CARPENTER (1992) noted that Chresmoda was in (1990). The fossils are preserved in a fine-grained fact, under his interpretation, an orthopteran and limestone, usually as simple impressions of the that it was often confused with Propygolampis (as animals, which are sometimes secondarily orange- in HANDLIRSCH’s synonymy list). Nonetheless, colored by iron-oxide. The three type specimens of CARPENTER (op. cit.) placed both Halometra and Sternarthron zitteli (and Halometra minor) are housed Sternarthron as junior synonyms of Propygolampis in the Bayerische Staatssammlung für Paläontologie and figured (CARPENTER 1992: 121) a slender-legged und Geologie in Munich (BSPGM), and are re- Propygolampis specimen of the form that OPPENHEIM described below: (op. cit.) called Halometra gigantean, and that HANDLIRSCH (incorrectly) called C. obscura.How- Re-description. – Specimen BSPGM labeled ever, following HANDLIRSCH (1906-1908), CARPEN- “413. 1870. VII. 45. Eichstaett” (Fig. 1.1): body 16 TER (1992) placed Propygolampis and its synonyms mm long; abdomen cone-shaped (length 9 mm) and (i.e. Sternarthron) in an uncertain family of the apically pointed, with an apparent short “terminal Phasmatodea, despite the fact that Propygolampis has filum” (length 1.5 mm), which is most probably either the overall appearance of a semiaquatic, gerromorph an artifact or an imprint of the cerci or an ovipositor; hemipteran. Also, misinterpreting HANDLIRSCH no details of head or body visible, except for traces of (1906-1908), CARPENTER (op. cit.) incorrectly stated large eyes, short antennae (length 2 mm), and maybe that HAASE placed Sternarthron in the Araneae. large biting mouthparts (mandibles); on both sides of As with HANDLIRSCH (1906-1808), CARPENTER’s the head there is a short appendage visible (length (op. cit.) work was initially overlooked and in his 5 mm left side, right side only 2.5 mm visible length) review of fossil arachnids, SELDEN (1993a)eschweizerbartxxx regarded that apparently originates close to the procoxa (left Sternarthron as a doubtful palpigrade while in The side) (most probably these appendages are palpi of Fossil Record 2,SELDEN (1993b) included Sternar- the mouthparts, or artifacts); only three pairs of legs thronidae as palpigrades, while reaffirming the need (profemur 15 mm, protibia only 4 mm, protarsus for restudy. In his review of the Solnhofen fauna, 10.5 mm; mesofemur 17.5 mm, mesotibia 8 mm, FRICKHINGER (1994) figured a Sternarthron [sic.] mesotarsus about 8 mm; metafemur 13 mm; metatibia specimen from a private collection, referring to it as 9 mm); a very distinct kink distal of profemur due to an arachnid; although in the accompanying English the very short protibia (forelegs); distance between text the German word ‘Spinnentiere’ (= arachnid) is procoxa and mesocoxa 2 mm, between mesocoxa and mistranslated as ‘spider’. Meanwhile, Sternarthron metacoxa only 1 mm (middle legs and hind legs was not included by CONDÉ (1996) in his review and closely parallel and directed backwards, and forelegs catalogue of palpigrades. SELDEN &DUNLOP (1998) widely separated from them and directed forwards, noted CARPENTER’s synonymy in their review of the exactly as in all adult chresmodids!). This was spe- arachnid orders, but miscited CARPENTER’s syste- cimen 413 of HAASE (1890: pl. 31, figs. 5-6), but his matics by calling Propygolampis a heteropteran drawings prove to be extremely imprecise when they insect. FRICKHINGER (1999) corrected the spelling are compared to the original fossil. This specimen to Sternarthron, while noting that the specimen he should be designated as the lectotype. figured previously (FRICKHINGER 1994: 159) was not Specimen BSPGM labeled “1964 XXIII 164 Sternarthron, but an unknown arachnid possibly Schernfeld” (plate and counterplate) (Fig. 1.2): body similar to opilionids. A further ‘Sternarthron’ from a 11 mm long; abdomen cone-shaped and apically private collection was figured by FRICKHINGER (1999) pointed, but without “terminal filum”; no details of who cites a pers. comm. by G. BECHLY (SMNS) that head or body visible, except faint traces of abdominal the fossil in question may be either a pycnogonid or segmentation and a pair of short antennae (length The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 359

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Fig. 2. 1: AS.VII.499, specimen studied and figured by GERMAR (1839) as Pygolampis gigantea (= Chresmoda obscura), Upper Jurassic (Tithonian) of Solnhofen (Germany); (BSPGM). 2: LP-94-IEI, Chresmoda aquatica (holotype), Lower Cretaceous (Barremian) of El Montsec (Spain). 3: LH-23300, Chresmoda sp., Lower Cretaceous (Barremian) of Las Hoyas (Spain). 4: NI 3a-b, Chresmoda libanica (holotype), Upper Cretaceous (Cenomanian) of Nammoura (Lebanon). 5: PIN 3149/1711, Saurophthirodes mongolicus (holotype) (Chresmoda mongolica comb. nov.), Lower Cretaceous of Gurvan- Ereniy-Nuru (Mongolia). 360 X. Delclòs et al. about 3 mm); only three pairs of legs (profemur 15 Cretaceous of Lingyuan in China, and C. libanica NEL mm; mesofemur 17.5 mm, mesotibia 8 mm, meso- et al., 2004 from the Cenomanian lithographic lime- tarsus 8.5 mm; metafemur 14 mm; metatibia 9 mm); a stones of Lebanon (Fig. 2.4). Copious specimens have distinct kink distal of profemur (forelegs); distance been discovered from the Upper Jurassic of Inner between procoxa and mesocoxa 2 mm, between meso- Mongolia (China), probably representing three dif- coxa and metacoxa only 1 mm. This specimen might ferent species (HUANG, pers. comm.). Other unde- eventually be the same as the original Dresden spe- scribed species are known from the Middle or Upper cimen of HAASE (1890: pl. 31, fig. 4), but this is very Jurassic of Bakhar in Mongolia (RASNITSYN & uncertain and just an assumption. QUICKE 2002), and from the Aptian – Albian of Specimen BSPGM labeled “AS I 822” (Fig.1.3): Santana do Carirí in Brazil (see below). Saurophthiro- body about 14 mm long; abdomen broad and without des mongolicus PONOMARENKO, 1986 from the Lower “terminal filum”; no details of head or body visible, Cretaceous of Gurvan-Ereniy-Nuru, in Mongolia (Fig. maybe except traces of a pair of short antennae; only 2.5) could be also included in Chresmoda. This shows three pairs of legs (profemur 15 mm, protibia only that the genus Chresmoda has almost a biostra- 4 mm, protarsus 10.5 mm; mesofemur 17.5 mm, tigraphic range Tithonian to Cenomanian with world- mesotibia 8 mm, mesotarsus about 8 mm; metafemur wide, mainly Laurasiatic, distribution. Sternarthron 13 mm; metatibia 9 mm); a very distinct kink distal of zitteli, are not Chelicerata – Pantopoda, but represent profemur due to the very short protibia (forelegs); the nymphal stage of C. obscura from the same distance between procoxa and mesocoxa 2 mm, locality. The holotype of C. obscura is missing today, between mesocoxa and metacoxa only 1 mm. This but the holotype of P.gigantea is housed in the Bayeri- specimen seems to be the type of Halometra minor sche Staatssammlung für Paläontologie und Geologie, OPPENHEIM (1888: pl. 31, fig. 4), and is most probably Munich, labelled AS.VII.499 (figured by GERMAR, pl. also the same as specimen 414 of HAASE (1890: pl. 22, fig. 8) (Fig. 2.1). 31, figs 1, 3). 3.1. A new species of Chresmoda from the Discussion. – CARPENTER (1992) already correctly Crato Formation limestones of Brazil recognized that the fossil arthropods from Solnhofen, known under the name Sternarthron zitteli, are not BECHLY (1998: 155) and BECHLY (1999: 9) briefly Chelicerata at all, but clearly represent theeschweizerbartxxx nymphal mentioned the discovery of chresmodid fossils from stage of Chresmoda obscura from the same locality. the Crato Formation for the first time. BECHLY et al. Our re-examination of the type specimens from the (2001: 55, fig. 44) discussed and figured a beautiful collection of the Museum in Munich (BSPGM no. fossil Chresmoda from the Crato limestones. This 1870 VII 45 and AS I 822) confirmed that the de- female specimen with ovipositor, which seems to scription and drawings of HAASE (1890) are com- be the best preserved specimen from this locality, is pletely erroneous, very freely drawn and “recon- housed with preliminary no. 0134 (old number H56) structed” (which also explains the dissimilarity to in the Staatliches Museum für Naturkunde Stuttgart in the drawing of OPPENHEIM, 1888), and incorrectly Germany. Another specimen from the American interpreted. These fossils clearly are hexapods with Museum of Natural History in New York was figured only three pairs of legs, and with leg structures that are by GRIMALDI &ENGEL (2005: fig. 7.5) (Figs. 3-5). very similar to adult Chresmoda (contra BECHLY Four further specimens have been studied by the 1999: 9). present authors. BECHLY (2007) discussed the chresmodids from the Crato Fm in more detail and provided a brief 3. The genus Chresmoda GERMAR, 1839 description, however without formally naming the The family Chresmodidae includes, besides Chresmo- new species. da obscura GERMAR, 1839 from the Tithonian litho- All six of the known specimens of Chresmoda graphic limestones of Solnhofen (Fig. 2.1), the similar neotropica n. sp. from Crato are adults and alate with C. aquatica MARTÍNEZ-DELCLÒS, 1989 from the long wings (wing length 27-28 mm). The body length Barremian lithographic limestones of Montsec (Fig. from head (without antennae) to abdomen (without 2.2) and Las Hoyas (Fig. 2.3) in Spain, the quite distal appendages) is about 21-25 mm, and the meso- different C. orientalis ESAKI, 1949 from the Lower femora are about 20-22 mm long. The head has large The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 361 globular compound eyes and large prognathous long; 2.26 mm wide); antennae inserted anteriorly and mandibles (similar to tiger beetles), and the antennae immediately adjacent to one another; scape swollen, are 9-11 mm long. Distinct, one-segmented cerci (3 approximately 1.5–2.0 times wider than pedicel, approxi- mately as wide as long; pedicel much smaller than scape, mm long) and a prominent ovipositor is visible in one approximately as wide as long; proximal flagellar articles as specimen. All other characters agree with the general wide as scape; subsequent flagellar articles decreasing in diagnosis of the lineage. width gradually toward apices of antennae; total number of Being surface striders on superficial fresh- or flagellar articles indeterminate as margins between articles are not discernable; mouthparts not preserved. Thorax brackish water (see also MARTILL et al. 2007) that moderately broader than head, approximately 1.6 times mainly fed on other insects that have been trapped on longer than wide (approximately 6.85 mm long; 4.24 mm the water surface, the chresmodids and hydrometrids wide), with prominent median keel; prothoracic basis- (NEL &POPOV 2000) most probably represented the ternum 2.26 mm long, 1.85 mm wide; mesothoracic only autochthonous aquatic insects in the paleohabitat basisternum 1.16 mm long at median keel, 3.08 mm wide; of the Crato lagoon. metathoracic basisternum approximately 3.29 mm long, 2.61 mm wide; pro–mesosternal intersegmental suture v-shaped towards posterior; meso-metasternal interseg- mental suture v-shaped towards anterior. Legs without Chresmoda neotropica ENGEL &HEADS,n.sp. spination; coxae robust, laterally directed; meso- and meta- Figs. 3-5 coxae markedly larger than procoxae; pro- and mesocoxae widely seperated; metacoxae apparently meeting at 1998 Chresmodidae. – BECHLY, p. 155. posterior margin of metathoracic sternum; protibia rela- 1999 Chresmodidae. – BECHLY,p.9. tively short (approximately 5.50 mm long), slightly 2001 Chresmoda. –BECHLY et al., p. 55, fig. 44. recurved, approximately 30 % of profemur length (18.92 2005 Chresmodidae. – GRIMALDI &ENGEL, p. 193, fig. mm); tarsi elongate, composed of between 8 and 12 tarsal 7.5. articles; meso- and metathoracic legs longer than protho- 2007 Chresmoda. –BECHLY, pp. 262-265, pl. 15d-e. racic legs; prothoracic leg 75% of mesothoracic leg, 80% of metathoracic leg. Only right forewing preserved; well Etymology: Specific epithet derived from ‘neotropical’; developed, approximately 27.80 mm long, at least 2.0 times in reference to the biogeographical provenance of the as long as abdomen (excluding ovipositor); venation largely species. indistinct; most veins simple and parallel–subparallel; SC long, simple, reaching wing margin before apex; sub- Type material: Holotype: Macropterous adult female, costal area proximally filled with dense, apically directed SMNS 66000-13 (old no. H56); housed in the Staatliches eschweizerbartxxx crossveins at least as far as midwing; RA and RP simple, Museum für Naturkunde, Stuttgart (Germany); Paratype: branching proximally of midwing; MA with two simple Macropterous adult AMNH SA-46446; housed in the branches; ?cubital area with dense pattern of crossveins American Museum of Natural History, New York (USA). forming minute cells, some of which lack pigmentation. Abdomen of similar width to thorax, total length (excluding Type locality: Chapada do Araripe, vicinity of Nova valvulae of ovipositor) approximately 13.88 mm long; Olinda, southern Ceará, north-east Brazil. abdomen apparently rotated somewhat postmortem to expose lateral surfaces of terga; subgenital plate triangular; Type horizon: Lower Cretaceous, upper Aptian, Nova valvulae of ovipositor elongate, forcep-like, approximately Olinda Member (limestones) of the Crato Formation 3.87 mm long; no serrations are discernable on the valvulae, (previously included in the Santana Formation sensu lato). though the distalmost part of the abdomen and the terminalia have been damaged somewhat during preparation Additional material: Incomplete adult (sex inde- of the specimen. terminate), UMB K25547 (Pl. 3, fig. 2); housed in the Ulster Museum, Belfast (Northern Ireland, UK); and a Description of paratype AMNH SA-46446 (Fig. specimen with no. G88 in coll. ms-fossil, Sulzbachtal, 5.1-5.4): Adult macropterous female, body form slender. Germany (featured in BECHLY 2007: pl. 15e). Head wider than long; antennae apparently positioned high on face, with interantennal (supraclypeal?) area Diagnosis: Interantennal (supraclypeal?) area swollen; slightly raised between antennal toruli, toruli separated thorax elongate, metacoxae meeting at posterior margin of by less than scape width; scape swollen, approximately metasternum, protibia relatively short (30% of profemur 1.5 times wider than pedicel or flagellar articles, length length), tibial spines absent, abdomen more elongate approximately 2.75 times width; pedicel about as wide as and narrow, margins subparallel for majority of their long; first flagellar article elongate, about three-fourths length. length of scape; total number of flagellar articles inde- terminate as antennae are incomplete; mandibles and other Description of holotype SMNS 66000-134 (Figs. mouthparts not discernable. Thorax only slightly broader 3.1, 5): Adult macropterous female preserved in ventral than head; thorax longer than wide, about 1.8 times longer aspect; body form slender. Head wider than long (3.21 mm than wide. Legs without spination; coxae widely separated 362 X. Delclòs et al. and laterally directed; protibia relatively short, 30% of 3.2.2. Legs profemur length (protibia 5.8 mm, profemur 19.2 mm); The legs are very long and narrow, covered with very tarsi elongate, with at least eight tarsal articles (total number cannot be ascertained in AMNH SA-46446); pretarsus (and short setae arranged in rows. These small setae likely its associated claws) either absent or modified into unit assisted in the ability of individuals to rest on the resembling a distalmost, apically-rounded tarsal article; water surface without penetrating, at least when being tarsi of preserved legs conspicuously surrounded or par- supported by floating water plants. As preserved, as tially covered by black crystalline material found no where likely also in life, the forelegs are projected forward, else in the matrix or on the compression. Wings fully developed, folded flat over abdomen when in repose, well while the mid and hind legs are projected backwards surpassing abdominal apex. Abdominal base broadly (Fig. 9.3). The general structure is similar in the three articulated with thorax and of equivalent width; abdominal legs but the tibial length is variable in respect to the terga wider than long (total number of abdominal segments femur for all species. The femora have different length not discernable), lateral margins subparallel, with terga respect to the tibia, which modifies the length of the becoming slightly and progressively narrower toward apex, quickly tapering in width in distalmost segments. Cerci and leg. For example, in adults of C. aquatica the pro- and ovipositor are not visible, and the thoracic nota are not well metafemora are equal in length while the mesofemora preserved. are longer; in C. obscura the meso- and metafemora are equal in length, and longer than the profemora. These differences are not so evident in the tibiae, but 3.2. The body structures of Chresmoda C. libanica have fore tibiae shorter than other studied species (nevertheless, all Lebanese specimens are Previous publications described accurately the struc- nymphs). Tarsi are usually difficult to see but in some tures of different species of the genus Chresmoda individuals it is possible to observe numerous small (HANDLIRSCH 1906-1908; ESAKI 1949; MARTÍNEZ- apical tarsomeres of progressively decreasing lengths DELCLÒS 1989; NEL et al. 2004; NEL et al. 2005). (Fig. 7.2). Presumably when these tarsomeres were Here we include and comment on the more interesting positioned on the water they created depressions in the features relevant to the phylogenetic placement of the surface without penetrating it owing to the backward family. curvature of the tarsi. The surface tension acted on the whole length of those tarsomeres that were in contact 3.2.1. Head with the water surface to support the weight of these Chresmoda has a prognathous head, completelyeschweizerbartxxx insects. All species studied have more than 40 tarso- covered by short, fine setae. The antennae are thick, meres that are superficially quite similar to antennal with 17 setaceous flagellar articles, with fine, short articles and have oblique, sigmoidal apices in both setae (in some specimens of C. obscura the antennal nymphal stages and adults (NEL el al. 2004), pre- setation is long and obscures the antennal segment- sumably acting as a locking mechanism to prevent the ation); the scape is long, the pedicel is 1/3 as long as complete distortion of the legs during rest (MARTÍNEZ- scape and the third segment is the longest. The total DELCLÒS 1991). length of the antenna is quite variable depending The procoxa – protrochanter articulation orients of the species and with respect to the body length, the protrochanter and more distal podites of the leg around 8 mm in C. aquatica, >12 mm in some spe- anteriorly. The mesocoxa – mesotrochanter articu- cimens of C. obscura, and 9-11 mm in Crato spe- lation typically orients the leg perpendicular with cimens; perhaps longer in C. orientalis with respect to respect to the length of the body but the midlegs are body length of other species. capable of being shifted widely forward and, pre- The presence of maxillary palpi, a short two- or sumably, backward as well (e.g., the preserved posi- three-segmented labial palpus without setae, and long tion of AMNH SA-46446 – Chresmoda neotropica n. and strong mandibles bearing a row of small obtuse sp. – has the midlegs projected forward nearly parallel teeth, and several short but sharp spines in the mouth with the forward-projecting forelegs). This same arti- suggest that the species of Chresmoda were predatory, culation in the hind legs projects the legs posteriorly. consuming small animals living at the water surface or The morphology of the coxa – trochanter articulation trapped by water surface tension (insects), or plank- suggests that their movement was oar-like. The mobi- tonic or nektonic animals when they came to the lity of the tibia with respect to the femur was weak surface. because in all studied Chresmoda the angle between femur and tibia was 100-110 degrees. The angle The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 363

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Fig.3.1:SMNS 66000-134; Chresmoda neotropica ENGEL &HEADS n. sp. (holotype), Lower Cretaceous (Aptian-Albian) of Santa Ana area (Brazil); housed in the Staatliches Museum für Naturkunde Stuttgart (Germany); general habitus of the species. 2: UMB K25547; Chresmoda neotropica ENGEL &HEADS n. sp.; housed in the Ulster Museum, Belfast (Northern Ireland, UK). 364 X. Delclòs et al.

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Fig.4.1:AMNH SA-46446; Chresmoda neotropica ENGEL &HEADS n. sp. (paratype), Lower Cretaceous (Aptian-Albian) of Santa Ana area (Brazil); American Museum of Natural History; general habitus of the macropterous species. 2: Head, basal part of the antenna and forelegs. 3: Distal part of the hind leg showing numerous small apical tarsomeres, and original pyrite cubes precipitate during the early diagenetic stages of fossilization. 4: Distal part of the femur and shorter tibia of the foreleg. The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 365

podites of the legs into antenna-like structures (see below). Current evidence indicates that this trans- formation probably took place during the Upper Triassic or Lower Jurassic, eventually becoming fixed in this lineage, and persisted for more than 65 Ma, disappearing along with the chresmodids during the Cenomanian (NEL et al. 2004).

4.3. Wings Hitherto now, individuals of Chresmoda were always found with their wings folded over the abdomen. PONOMARENKO (1985) attributed this taphonomic feature to their special mode of life at the water sur- face. Some individuals of C. obscura with unfolded wings was found in Solnhofen (i.e. JM.1958a and JM.1964.2, both from the Juras-Museum in Eichstätt), although its degree of preservation prevented detailed study of the wings. Usually terrestrial insects that have flown to water and have been trapped by the surface tension of the water die by asphyxiation and unfold their wings (MARTÍNEZ-DELCLÒS et al. 2004). One exception is the wings of orthopterans that are typi- cally folded at rest owing to the nature of their articu- lation. Because chresmodids are broadly related to orthopterans, they may have had a similar form of articulation that promoted their post-mortem retention in a folded position. It was not possible to accurately compare the methods of articulation between chres- eschweizerbartxxx modids and other lineages of insects, most notably the Orthoptera. Two new specimens of Chresmoda sp. with pre- served fore- and hind wings (LC-1123-IEI, Barremian of El Montsec, and LH-26.546 of Las Hoyas) were recently discovered in Spain. The pattern of venation of LC-1123-IEI (Fig. 8.1-8.3) is nearly identical to Fig.5.1:SMNS 66000-134; Chresmoda neotropica ENGEL those of the Mesozoic “stick insect-like” taxa of the &HEADS n. sp.; camera lucida drawing of the holotype. order “Phasmatodea” sensu GOROCHOV (1994). It has bst: basisternite, C: cubital vein, fg: flagellomere, mcx: mesocoxa, mtcx: metacoxa, mtr: mesotrochanter, mttr: elongate wings (Fig. 8.3), straight radial, median and metatrochanter, p: pedicel, pcx: procoxa, scp: scape, s: cubital veins, all parallel, with numerous straight sternite, SC: subcostal vein, sgp: subgenital plate, t: tergite. cross-veins between them; few elongate intercalary longitudinal veins; ScP ending on the wing margin not far from the wing apex; MA with two simple branches; RA, RP, MP, CuA, and CuP simple; an between the tibia and tarsus was more constant, apical anastomosis between the median veins in the around 180 degrees (Fig. 4.4). We consider that all hind wing; a broad anal area in the forewing; and a skating effort was done by the basal part of the legs very large and broad vannus in the hind wing. The (coxa through tibia). pattern of venation of LH-26.546 (Fig. 6.2-6.4) is This dramatic specialization, unique among aquatic even more interesting as its forewing base shows for Hexapoda, may be perhaps the product of a homeotic the first time the exact structure of the median and mutation affecting genes that participate in leg deve- cubital veins. Its concave CuP has a very short con- lopment, resulting in the transformation of the distal cave anterior branch CuPa that extends towards the 366 X. Delclòs et al.

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Fig. 6. LH-26.546; Chresmoda aquatica, Lower Cretaceous (Barremian) of Las Hoyas. 1: General habitus of the specimen. 2: Base of the forewing. 3: Camera lucida drawing of the forewing. 4: Camera lucida drawing of a detail of the base of the same forewing. base of the convex M+CuA. This last vein separates parallel branches, RP separates from RA in a very at the same point into a convex anterior MA and a distal position, and the preserved part of RA is simple convex posterior branch MP+CuA. MP separates and straight. Nearly all of the longitudinal veins are from CuA significantly more distally. This pattern is parallel and straight (see Figs. 5.1, 6.3-6.4, and 8.3). typical of polyneopteran insects of the Archaeortho- Some specimens from Solnhofen also have long ptera (BÉTHOUX &NEL 2002). Veins CuPb, CuA and wings, overlapping the abdomen, similar to the spe- MP are straight and simple, MA has three long and cimen LC-1123-IEI from El Montsec, and are thus The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 367

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Fig. 7. Chresmoda aquatica, Lower Cretaceous of Las Hoyas. 1: LH-13826, brachypterous female specimen. 2: distal part of the mid and hind legs; see the numerous small apical tarsomeres that compound the leg. 3: LH-13574, apterous male specimen preserved in organic rich mudstones. 4: LH-92AM102, brachypterous female preserved in non-organic rich mudstone level. 5: MCCM-LH 18023, nymph that shows the numerous small apical tarsomeres. 6: MCCM-LH 18024, isolated forewing highly sclerotized, possibly from a brachypterous female specimen. All specimens housed in the Museo de las Ciencias de Castilla-La Mancha in Cuenca (Spain). 368 X. Delclòs et al.

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Fig. 8. LC-1123-IEI, Chresmoda aquatica, Lower Cretaceous of El Montsec. La Cabrua outcrop. 1: Right forewing. 2: Right hind wing with a developed vanus. 3: Camera lucida from the fore- and hind wing of the specimen, showing the vein pattern distribution. Housed in the Institut d’Estudis Ilerdencs in Lleida (Spain).

macropterous. But some specimens from Solnhofen, All adult females (large specimens with ovipositor) Las Hoyas, and El Montsec have distinctly shorter were winged (Fig. 9.1), whereas adult males (large wings relative to the length of the abdomen, just specimens without ovipositor), and nymphs (smaller scarcely overlapping the abdomen, with the forewing specimens, with or without ovipositor) were apterous hairy and probably heavily sclerotized (Fig. 7.1, 7.4). (Figs.3.2,7.5). This contrasts with the usual secondary The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 369

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Fig. 9. LP-94-IEI. Chresmoda aquatica, holotype; brachypterous specimen, Lower Cretaceous of El Montsec. 1: distal part of the abdomen showing the ovipositor and cerci. 2: reconstruction of the distal part of the abdomen; see the unsegmented cerci and serrate ovipositor. 3: apices of the fore- (right) mid- and hind femur (left) in connection with the tibia; please note the blocking structure between femur and tibia that restricts anterior or posterior movements of the legs (forelegs cannot be moved backwards, while mid- and hind legs cannot be moved forwards). The arrow indicates the anterior orientation in direction of the head. Housed in the Institut d’Estudis Ilerdencs in Lleida (Spain).

apterous females and alate males, in some ortho- segmented cerci (Fig. 9.1.-9.2). The ovipositor is long pteroid and phasmatodean species. Perhaps the well- with two valvulae and an “orthopteroid” gonoplac. developed wings of Chresmoda females increase their The valvulae and gonoplac bear serrations (Fig. 9.2) possibility to escape from other predaceous insects, or that might be correlated with endophytic egg-laying, to disperse to new aquatic environments during dry perhaps in floating plants (NEL et al. 2004) or into seasons. helophytic ones, such as Frenelopsis. The ovipositor sheath is also present in young nymphs from Lebanon and Las Hoyas. The brachypterous females have 4.4. Abdomen shorter and stronger abdomens than macropterous The abdomen seems to be 8- or 9-segmented and specimens (Fig. 7.4). The presence of wings in C. distally possesses two small, sharp, pilose, one- orientalis suggests, in relation with other species of 370 X. Delclòs et al.

Chresmoda, that it is a female, but no ovipositor is dea]. CAMERON et al. (2006) proposed a relationship described nor observed. The presence of cerci, among [Grylloblattodea (Mantophasmatodea + Timemato- other characters, definitively excludes placement dea)], but neither included the Phasmatodea s.str. (= among the Paraneoptera and historical attributions to Euphasmatodea or Euphasmida) nor the Embioptera Hemiptera can be accordingly dismissed. in their analysis. A paraphyly of Phasmatodea (= Timematodea + Euphasmatodea) is clearly contra- 4. Phylogenetic relationships dicted by the results of WHITING et al. (2003), and TERRY &WHITING (2005). 4.1. Wing venation The Chresmoda hind wing LC-1123-IEI from El In the last years significant progress was achieved Montsec (Fig. 8.2-8.3) shows some similarities with in the basal phylogeny of Recent Phasmatodea those of the Cenozoic and Recent Phasmatodea (true (BRADLER 2003; KLUG &BRADLER 2006). However, stick- and leaf-insects), especially in the expanded the phylogenetic position of the crown-group Phasma- anal area and mains veins simple, straight and parallel. todea is still very controversial. The order has been The forewings of the majority of Recent Phasmatodea considered as the sister-group of Dermaptera (KAMP are reduced or absent, with a strongly sclerotized 1973), Embioptera (RÄHLE 1970; TILGNER 2001; wing base. Such a character state was recently also WHITING et al. 2003; ZOMPRO 2004; KJER 2006), described from an Early Tertiary leaf insect by Grylloblattodea + Dictyoptera (MAEKAWA et al. 1999, WEDMANN et al. (2007). If this reduced state of the who did not include Embioptera in their analysis), wings is a groundplan feature of Phasmatodea is still a Dictyoptera (BEUTEL &GORB 2001), Orthoptera (ALI matter of debate (WHITING et al. 2003; TRUEMAN et al. &DARLING 1998; FLOOK et al. 1999; WHEELER et al. 2004; WHITING &WHITING 2004). 2001; GOROCHOV 2001; BEUTEL &GORB 2006), as TILGNER (2001) discussed the position of the “pre- an ‘orthopteroid’ order (KUKALOVÁ-PECK 1991; Tertiary Phasmatodea” attributed to the “Phasmo- GRIMALDI 2001; GRIMALDI &ENGEL 2005), as sister ptera” sensu GOROCHOV (1994) [Susumanioidea + to all other Polyneoptera (DALLAI et al. 2005), or as a Prochresmodoidea + Aeroplanoidea + Xiphopteroi- paraphyletic group, viz. one group ‘Timematodea’ dea], and concluded that they are probably not related sister to the Embioptera, together representing the to the Recent and Cenozoic Phasmatodea. As no sister of the Plecoptera, and the whole set [(Time- phylogenetic analysis of the “Phasmoptera” or “pre- matodea + Embioptera) + Plecoptera], as theeschweizerbartxxx sister- Tertiary Phasmatodea” is available, the few characters group of ‘Phasmatodea’ s. str. (ZOMPRO 2005). that could be potentially used are not polarized (but DALLAI et al. (2003, 2007) studied the sperm ultra- see WILLMANN 2003). At least some representatives structure and failed to support a sister-group relation- (Orephasma REN, 1997; Aeroplana TILLYARD, 1918) ship of Phasmatodea with Embioptera. DALLAI et al. of these “pre-Tertiary Phasmatodea” have specialized (2005) proposed a polytomic [Mantophasmatodea + structures of the forewing cubito-median veins: a Mantodea + (modern) Orthoptera + (modern) Gryllo- basal fusion of convex CuA with M into a common blattodea] supported by a single, unique apomorphy, stem, CuA re-emerging from M a short distance from i.e., ‘three connecting bands’ in sperm structure. wing base, and separating from a convex MA and These authors indicated that the situation in Ortho- a concave MP, and a concave CuP divided into ptera is polymorphic going from total absence to one, branches, one of them reaching M+CuA, or two, or three such bands, and that they a priori chose MP+CuA. These structures are putative synapo- the above state as the ancestral condition for Ortho- morphies of the “orthopteroid” lineage. RAGGE (1955) ptera because it is ‘the most frequent one in the did not observe them in Recent Phasmatodea. These group’. This method of estimation of an ‘ancestral structures are as difficult to observe in the Recent state’ is utterly unfounded, without merit, and should Phasmatodea as in the modern Orthoptera owing to be rejected. Analysis with Orthoptera coded as the strong sclerotization of the wing bases and basal polymorphic for this character leads to a clearly much displacement of the stems of the median and cubital less resolved tree. As such, this analysis must be veins. But they are very clearly visible in the Palaeo- entirely reconsidered. Like ZOMPRO (2004), KJER et zoic and Mesozoic Orthoptera and in some Mesozoic al. (2006) placed the modern Phasmatodea as para- “Phasmatodea”. It would, however, be necessary to phyletic group near Embioptera, and together as re-examine all pre-Tertiary “Phasmatodea” to de- sister-group of [Grylloblattodea + Mantophasmato- termine their exact patterns of venation. Some of The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 371 these insects may well have been related to modern present in basal Recent mantid taxa like Chaetessa, phasmatodeans while others were unplaced members Mantoida, and Metallyticus. As in Chresmoda that of Archaeorthoptera (such as Orephasma and Aero- had more prominent stiff, stout hairs (or spines) on the plana). rear of the legs, mantids have evident spines in the Fortunately, the area of the branches of CuP and same position. Mantis tegmina are more or less hard base of M+CuA is preserved in the forewing of the and coriaceous (Fig. 7.6). Some brachypterous species Chresmoda LH-26.546 from Las Hoyas (Fig. 6.3-6.4). of Chresmoda had coriaceous tegmina (with more or It corresponds exactly to the venational pattern of the less evident venation), whereas in other individuals Clade Archaeorthoptera [= Orthopterida] and, there- the wings are completely membranous. The abdomen fore, supports the attribution of Chresmodidae to the of mantises is wider in females than in males, while in “orthopteroid” rather than to the modern “phasma- chresmodids the abdomen is morphologically dif- todean” lineage. ferent depending on whether the individual was apterous, brachypterous, or fully winged. Female Comparison of Chresmoda with the Meso- mantises have largely vestigial valvulae owing to the zoic “Phasmatodea” (Holophasmatodea). – deposition of eggs in oothecae. All Cretaceous stem- The great majority of Mesozoic “Phasmatodea” group mantises are obvious Mantodea (GRIMALDI 2003) and bear no relationship to Chresmodidae, (Holophasmatodea sensu GRIMALDI &ENGEL, 2005) are known only from isolated forewings. Chresmoda which also lack predatorial forelegs, contrary to all known fossil and extant mantises. differs from the Susumaniidae GOROCHOV, 1988 in its simple RP,branching from R in the middle of the wing (GOROCHOV 1988, 2000). GOROCHOV (1994) charac- Comparison with Phasmatodea. – Phasma- terized the Xyphopteroidea SHAROV, 1968 by the todea and Chresmoda are prognathous insects with pectinate MP in the forewing, quite unlike that seen in monofiliform to filiform antennae. They have a short Chresmoda. The Necrophasmatidae also have a prothorax. Many species are apterous or brachy- posteriorly pectinate MP (SHAROV 1968). The Aero- pterous. Phasmatodeans have the posterior wings planoidea TILLYARD, 1918 have RP, MP, or CuA with with a sclerotized anterior region and small, widely- several branches, unlike that state seen in Chresmoda. separated coxae, as in Chresmoda. The trochanter of Within the Prochresmodoidea VISHNIAKOVA, 1980, the Euphasmatodea is considerably smaller than of Chres-

eschweizerbartxxx Permophasmatidae GOROCHOV, 1992 has RP forked modidae, usually fused to the femur, which can be and a zigzagged intercalary vein between RA and RP autotomized at its boundary (see also WEDMANN et al. (GOROCHOV 1994). Chresmoda shows the greatest 2007). The femora and tibiae are long in both groups. similarities with Prochresmodidae VISHNIAKOVA, Nonetheless, in Chresmoda a notable reduction of the 1980 in its simple RP, MP, and CuA, and forked MA tibiae is present. Phasmatodeans have pentamerous (NEL et al. 2004). tarsi (the plesiomorphic condition for Insecta), but some regenerate legs can have tetramerous tarsi. In male chresmodids there are no sclerotized structures 4.2. Other characters compared visible that resemble the vomer of Recent Phasma- todea. Contrary to chresmodids, Recent female Some of the characters listed below are difficult to phasmatodeans have a reduced ovipositor composed polarize. The pattern of forewing venation remains the of the standard orthopteroid form (i.e., two valvulae best evidence supporting an attribution of Chresmoda and a gonoplac that assumes a valve-like form and to the Archaeorthoptera. function). In the huge majority of phasmatodean species, the ovipositor is concealed by the abdominal Comparison with Mantodea. – Mantises are sternum 8 (operculum), thus being internal (see orthognathous whereas chresmodids were pro- BRADLER, 2003: fig. 16.3). According to BEDFORD gnathous. In the antennae of mantises, the third (1978), phasmatodeans have 5-7 nymphal instars, antennomere is the longer, not observed in Chresmo- according to BRADLER (2003) 4-8 instars, with fe- da, which is equal in length. In most mantises the males usually having more (!) instars than conspecific pronotum is elongate, with a central sulcus, unlike the males. more or less rounded pronotum of nymphs and adults The number of nymphal instars in Chresmoda is of Chresmoda. However, a short pronotum is still unknown. Some species of phasmatodeans are parthe- 372 X. Delclòs et al. nogenetic (e.g., Bacillus, Clonopis, etc.) with many pterida]. Thus, if the Mantophasmatodea were actually females and few to no males. It might have been the related to the Orthoptera sensu stricto, they would same for Chresmoda, as we found in the fossil record presumably be more derived than many other Archae- more female than male specimens, but this could also orthoptera [= Orthopterida]. The Mantophasmatodea be due to other reasons. are increasingly understood to be the sister taxon WEDMANN et al. (2007) described a fossil leaf to Grylloblattodea [in Notoptera = Grylloblattodea + insect including the temporal sequence of character Mantophasmatodea] (ENGEL &GRIMALDI 2004; evolution that led to the extant crown-group. Charac- GRIMALDI &ENGEL 2005; TERRY &WHITING 2005; ters such as the basally curved fore femora, which are ARILLO &ENGEL 2006; KJER et al. 2006), although adaptations for camouflage and catalepsy at daylight the definitive placement of the group remains contro- in Recent Phasmatodea, are obviously not developed versial (CAMERON et al. 2006). The absence of salta- in Chresmodidae, suggesting that these insects were torial legs in Chresmoda is certainly related to its not nocturnal. mode of life and indicates nothing about its phylo- genetic affinities as this could merely be autapo- Comparison with Orthoptera. – Chresmoda morphic. shares with the Orthoptera their long, tripartite ovi- positor (composed of two valvulae and the valve-like 4.3. Other arthropods with multisegmented gonoplac). The relatively short antenna of Chresmoda tarsi could be an adaptation to aquatic environments. In Orthoptera the scape and pedicel are larger than the Until now, no other hexapod has been known to have flagellar articles, whereas the pedicel is the shortest super-multiarticulate legs, with an extreme sub- article in Chresmoda. Macropterous, brachypterous, division of an individual podite as is seen in Chresmo- or apterous taxa are known in Orthoptera, as in da (NEL et al. 2004). Hexapods have had since the Chresmoda. Orthopteran ovipositors show denticulate Late Carboniferous, and likely since the Devonian, inferior valvulae, as in Chresmoda. In addition, the five or fewer tarsal articles. robust metacoxae meeting at the midline of the Chresmoda have the last two tarsomeres of all legs sternum in Chresmoda neotropica are atypical for subdivided into more than 40 minute articles. Some Orthoptera, which usually have small, well-separated Recent and fossil groups of terrestrial arthropods metacoxae. eschweizerbartxxx show multi-segmented legs. Usually these modifi- The Chresmoda also have one-segmented cerci. cations are only present in the forelegs, and are related Monomerous cerci are present in Orthoptera and most to sensorial functions. Such is the case in arachnids Recent Phasmatodea, while the cerci tend to be poly- like Amblypygi, Uropygi and Schizomida, in which merous in most dictyopterans. Although ZOMPRO the first leg ends in a subdivided, antenniform tarsus (2005) proposed unsegmented cerci as a synapo- with a sensorial function. This is particularly apparent morphy of his ‘Orthopteriformia’ (= Mantophasma- in the slender forelegs of amblypygids. Fossils of todea + Orthoptera), the exact phylogenetic value of Amblypygi and Uropygi are found in the Carboni- this character remains uncertain because it is widely ferous and Cretaceous; Schizomida in the Pliocene homoplastic across Polyneoptera (TILGNER et al. (e.g., see SELDEN &DUNLOP 1998). Forelegs of 1999; TILGNER 2001; GRIMALDI &ENGEL 2005). The Palpigradi have their first pair of legs multi-articu- female terminalia of the Phasmatodea are of a typical lated with a sensorial function. In palpigrades the first orthopteroid type, as is also the case for Gryllo- pair of legs never touches the ground, in contrast with blattodea (DEUVE 2001), and thus apparently of the arachnid groups mentioned above. Fossils of this little interest for our purposes. Palaeozoic and some group have been found but are exceptionally rare Mesozoic stem-group dictyopterans (the so-called (ROWLAND &SISSOM 1980). Opiliones have in all four “roachoids”: GRIMALDI 1997; GRIMALDI &ENGEL pair of legs a multi-segmented tarsus (up to 100, that 2005) had long, external ovipositors similar to that permit tarsi to roll around plant axes) and with distal observed in Chresmoda. ZOMPRO (2005) also pro- claws on pretarsi, and their functionality are only for posed the saltatorial hind legs as a synapomorphy of walking; the earliest harvestmen are found in the the ‘Orthopteriformia’, but BÉTHOUX &NEL (2002) Devonian of Rhynie, but mainly in Tertiary ambers. demonstrated that saltatorial legs are not present in All of these groups of terrestrial arachnids have a the most basal lineages of Archaeorthoptera [= Ortho- scarce fossil record. No continental aquatic arthropod The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 373 has either analogous or homologous leg modifi- podite is separated from the next by a more or less cations. flexible joint composed of arthrodial membrane (the trochanter-femur joint is frequently immobilized) and controlled by musculature originating inside of the 5. Palaeobiology of Chresmoda body or within the preceding podite (the subdivisions Prior to this work it was supposed that the number of of the tarsus, the tarsomeres, are not true podites and tarsomeres increased along the evolution of the genus there is no intrinsic musculature operating them; Chresmoda during the Mesozoic. Chresmoda was muscles only move the entire tarsus as a unit), with the presumed to have pentamerous (C. obscura in exception of the pretarsus which is controlled by PONOMARENKO 1985) or tetramerous tarsi (C. aqua- muscles that extend back through the tarsus to the tica in MARTÍNEZ-DELCLÒS 1989). MARTÍNEZ- apex of the tibia or the femoral base. Most of the DELCLÒS (1991), studying new material from the studies on segmentation and development in insects Barremian of Spain, proposed that the last tarsal have been undertaken on the fruit fly Drosophila. article was subdivided into almost eight subsegments Segmentation is a developmental mechanism common that increased in length apically. A pretarsus with to a large number of animal lineages. Segmentation claws could not be discerned and instead in C. aqua- subdivides a tissue into a series of repeating units, tica the apex of the leg is apically rounded, either whereupon each basic unit can then be further ela- terminating with the last, subdivided tarsal article (and borated upon during development (RAUSKOLB 2001). thereby entirely lacking the pretarsal podite which The podites of the legs are not segments in the sense bears the claws) or in an apically-rounded pretarsal of body segmentation (i.e., developmental meta- podite with ungues lost. meres) but are instead subdivisions of metamere NEL et al. (2004) studied a then new species of appendages. Nonetheless, the development and con- Chresmoda from the Cenomanian of Lebanon. That trol of the formation of these subdivided appendages species had the three basal tarsomeres unsegmented has been studied in detail only within model orga- but the distal parts of the legs are divided into more nisms such as D. melanogaster, with cursorial ex- than 40 exceptionally small subtarsal articles, de- aminations for others like Gryllus, Oncopeltus,or creasing apically in length in the mid and hind legs, Apis, and the development of most hexapods remains and with 15 such units in the forelegs. Subsequent utterly unstudied. Although the molecular basis of to their work new material from Lebanon,eschweizerbartxxx new segmentation and regional growth during morpho- specimens from the Barremian of Las Hoyas and El genesis of insect legs is poorly understood, early steps Montsec (housed in Spanish museums), and from the in insect leg development have been elucidated, and Tithonian of Solnhofen (housed in the Natural History some key genes involved in leg segmentation have Museum, London and the Muséum national d’Histoire been identified (BISHOP et al. 1999; DE CELIS et al. Naturelle, Paris) demonstrate that both C. aquatica 1998; RAUSKOLB &IRVINE 1999). It is now well and C. obscura, respectively have more than 40 distal established that four-jointed (fj) interacts with dachs subtarsal articles (DA and XD pers. obs.). The same is (d), abelson (abl) and enabled (ena) and feeds back true for C. neotropica n. sp., reported herein from the onto the Notch (N) pathway involving the N ligands Crato Formation. This extremely rare segmentation of Serrate (Ser) and Delta (Dl) to affect growth and tarsi is observed also in minute nymphs from Las segmentation in the Drosophila leg (BUCKLES et al. Hoyas. 2001). Concerning tarsal segmentation, most of the Excluding Chresmodidae, the legs of insects are tarsus of insect legs derives from cells expressing typically composed of six podites (the next to last of Distal less (DLL). The studies of RAUSKOLB (2001) which, the tarsus, is divided into a maximum of five demonstrated that DLL represses N ligand expression articles, or tarsomeres, giving a total apparent number and that spineless-aristapedia (ss) regulates the of nine units in the leg) from proximal (closest to the expression of bric-à-brac (bab) which is also required body wall) to distal (tip of the leg) they are the coxa, for the subdivision of the tarsus into individual sub- trochanter, femur, tibia, tarsal articles 1-5, and the segments. HERKE et al. (2005) confirmed that in pretarsus (FRISTROM &CHIHARA 1978; GRIMALDI & Oncopeltus fasciatus, tiptop is a selector gene that is ENGEL 2005). The pretarsus bears the claws which are required for the segmentation of the distal leg and is frequently and quite erroneously called “tarsal claws” also required to switch appendage development from but are, in fact, “pretarsal claws” or “ungues”. Each antenna to leg. For some authors antennae and legs are 374 X. Delclòs et al. considered to be serially homologous structures as meable and pilose). Although they possessed different both are metameric appendages (GRIMALDI &ENGEL apical leg structures, it is possible that like Recent 2005) and because they can be interconverted through gerrids Chresmoda made whirlpools that affected the action of homeotic genes (EMERALD et al. 2003). levels in the water surface and with backward move- One could conclude that the leg and tarsal subseg- ments propelled the insect forward (HU et al. 2004). mentation is a complex, polygenic trait. For most of Gerrids transfer quantity of movement to the fluid the above cited genes induced mutations generating surface with the help of hemispheric whirlpools loss of function result in a reduction of tarsomere produced by their propulsive mid and hind legs. It is number, and yet, it remains unclear how a repeating possible that Chresmoda could have developed similar segmental pattern is generated during leg deve- hemispheric whirlpools (U-shaped) but with all three lopment, but one could imagine that over-expression pairs of legs. HU et al. (2003) showed that for 342 of these same genes could induce a multiplication of species of water strider the length of the leg in contact tarsomere number. Of course this kind of increase in with the water surface increases relative to the body tarsomere number must be followed by subsequent weight and thus the force due to surface tension modification of associated tissues requiring the action increases with the force due to body weight. It was of additional structural genes. Since the tarsus lacks apparently the same in Chresmoda, as the largest intrinsic muscles, this would not require modified species, C. obscura, has legs proportionally longer musculature but would require a great elongation of than in other species. the pretarsal depressor (the pretarsus lacks a levator Chresmoda aquatica, C. mongolica comb. nov., muscle) or unguitractor tendon, assuming that the and C. orientalis lived in lacustrine environments, pretarsus was retained in Chresmoda in some modi- while C. obscura and C. libanica apparently lived in fied form (perhaps resembling an apicalmost tarso- brackish-marine waters. The new species from Crato mere), something we cannot confirm. It is possible (Brazil) lived in a saline lagoon (GRIMALDI &ENGEL that during the evolution of the lineage that gave rise 2005; NEL et al. 2005). Data concerning the sensi- to Chresmodidae there were mutations involving tivity of Chresmoda species to increased salinity homeotic genes for leg segmentation that resulted in levels of other forms of environmental changes are not increasing the tarsomere number in the family. This available. In general, Gerridae are moderately tolerant mutation was fixed and successful for at least 65 to salinity levels (GOODERHAM &TSYRLIN 2002; Million years. However, we must also noteeschweizerbartxxx that CHESSMAN 2003). Pseudomorphs of salt cubes are any dramatic anatomical change (i.e., most synapo- usually found in the laminated limestones of Crato morphies) represents mutations of one form or (MARTINÉZ-DELCLÒS et al. 2004: 42, fig. 7C) but another that become fixed and successful in the clades perhaps the lagoon of Crato was solely saline in its that they define. While we have highlighted the bottom or in its water table (see also MARTILL et al. peculiar leg morphology of Chresmoda, other muta- 2007), and the water surface itself was fresh such that tions across the insects gave rise to the eventual origin chresmodids could skate there upon. of wings, various wing-folding mechanisms, the Hitherto now, it seems that the more ancient and modification of wings into a variety of structures earliest species of Chresmoda lived in brackish or of varying functions (e.g., halteres, elytra, blood marine water surfaces, while the Early Cretaceous sinuses), among innumerable others, and several of species are found only in lacustrine environments. them, but not all, has led to a successful diversifi- Owing to the absence of an accurate cladistic phylo- cation of hexapods. genetic position of the genus Chresmoda solidly The densely pilose tarsi in Chresmoda seem to be placing the group among the orthopterids (although a specialization for skating across water surfaces certainly more closely related to the Orthoptera than (MARTÍNEZ-DELCLÒS 1991; NEL et al. 2004) but the Phasmatodea), it is impossible to know when surely not in the same manner as in the modern the homeotic mutation that promoted this hyper- Gerridae (Hemiptera: Heteroptera) owing to their specialized leg structure took place. Nevertheless, we overall different leg structures. The pressure of a leg can hypothesize that this transformation likely took produces a trough-shaped depression (meniscus) on place during the Late Triassic or Early Jurassic, the water surface, and the surface tension of the water favoured by the decreasing of water density as a result easily supports the weight of the insect. The weight of a high global temperature during the Triassic (Fig. of Chresmoda was probably held up by the surface 10). curvature that developed around the legs (imper- The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 375

eschweizerbartxxx

Fig. 10. Period of appearance of chesmodid species in relation with the atmospheric CO2 average, and global climatic situation. 1: Graphic showing the details of CO2 proxy data set (Four-point running averages for individual proxies: Paleosols, Phytophacton, Stomata, and GEOCARB III), and combined atmospheric CO2 concentration record as determined from proxies (Grey curve represents average values in 10 My (after ROYER et al. 2004). 2: Average of the global temperature, and distribution of the “Ice House” – “Green House” situation in the Earth between Today since the Permian (after Scotese Paleomap 2003 web). 3: Distribution of the Chresmoda species during the Jurassic-Cretaceous. 376 X. Delclòs et al.

During the Late Jurassic the global climate changed Brazil (NEL &POPOV 2000), latest Albian amber of due to the break up of Pangea. The oldest studied Burma (ANDERSEN &GRIMALDI 2001; GRIMALDI et species of Chresmoda comes from the Tithonian of al. 2002; CRUICKSHANK &KO 2003), or in the Solnhofen, at 40º of latitude, living under a warm Cenomanian amber of Charente-Maritime, France temperate climate (SELLWOOD et al. 2000), but during (PERRICHOT et al. 2005). From the available evidence an otherwise global icehouse period (BERNER 1998; it does not appear as though gerrids and/or hydro- VEIZER et al. 2000; LABANDEIRA 2006). The climate metrids were competitors excluding chresmodids was more stable and uniform than today and in low from the same ecological niche. latitudes no evidence of tropical rainforest exists (REES et al. 2000, 2004). Except for C. libanica, all 6. Conclusions species of Chresmoda are found in rocks formed under warm temperate climates with moderate humi- The Jurassic-Cretaceous genus Chresmoda is mor- dity and driven by monsoonal seasons in low-mid phological and phylogenetical restudied. After critical latitudes (TAJIKA 1998). BARRON et al. (1989, fig. 13) revision we include the genus Sternarthron from the suggest that a globally warm climate meant high Upper Jurassic of Solnhofen which was previously precipitation and evaporation rates, predicting high placed among the arachnids, principally the palpi- seasonal rainfall focused on the northern and southern grades. borders of Tethys, where diverse species of Chresmo- We also review the principal morphological charac- da lived. The youngest species, C. libanica, is found ters of Chresmoda in order to elucidate their phylo- in the Cenomanian (under warm greenhouse global genetic relationships with other groups of ortho- climate). Its extinction is difficult to explain but we pteroids and arthropods, and we describe the new can hypothesize that it was perhaps due to the extreme species Chresmoda neotropica ENGEL &HEADS n. sp. global warming during the Turonian (BICE et al. 2003; from the Aptian-Albian lithographic limestones of the ROYER et al. 2004; JENKYNS et al. 2004). This is not Crato Formation in Brazil. The new specimens of a wholly satisfactory hypothesis because the global Chresmoda found in the Barremian of Spain with well climate changed considerably between the Tithonian preserved fore- and hind wing venation permit us to and the Cenomanian, evolving in middle latitudes determine that the venational pattern follows that of from warm temperate to very warm intertropical. the Archaeorthoptera (= Orthopterida) and presently The largest known species of Chresmodaeschweizerbartxxx (C. supports attribution of Chresmodidae to the “ortho- obscura) lived during the Tithonian, when the water pteroid” rather than to the modern “phasmatodean” surface temperature was low, with a consequent lineage. increase in water density. During the late Early One of the more impressive characters is the Cretaceous and the Late Cretaceous, a “greenhouse specialized structured legs adapted for skating across climate” was globally established (GALE 2001, water surfaces. During the evolution of the Chresmo- LARSON et al. 1993). In order to float in lacustrine da lineage a mutation involving homeotic genes environments, Chresmoda species were “obliged” to promote leg segmentation that resulted in increasing decrease their dimensions. During the Cenomanian it the tarsomere number in the family. This mutation was necessary for species of Chresmoda to occupy was fixed and successful for at least 65 Ma. A new brackish or marine environments with a higher water hypothesis about their spatial and environmental density in order to float. Perhaps Chresmoda were distribution thorough time based on the climate also aided by the large development of shelf areas in evolution is proposed. Their extinction during the marine environments caused by a global transgression Late Cretaceous remains unresolved, but likely was (HALLAM 1992). The Cenomanian was a period of not a result of competition for “ecospace” with the overall sea-level rise. semi-aquatic heteropterans. It is not possible to correlate the disappearance of Chresmoda with the origin and rise of other water- Acknowledgements striding insects, such as species of Gerridae or Hydro- metridae, since these families of aquatic heteropterans This research was made possible thanks to the active help of ALAIN WALLER,CHRISTIANE AMEDEGNATO, and SIMON were already established during the Cretaceous [e.g., POULAIN (Entomology, MNHN, Paris). We thank BILL Albian amber of Peñacerrada, Spain (DELCLÒS et al. SHEAR and ANDY ROSS for valuable discussions and DAV E 2007), Aptian-Albian laminate limestones of Crato, UNWIN for comments on the Solnhofen locality. Parti- The enigmatic Mesozoic insect taxon Chresmodidae from the Lower Cretaceous of Brazil 377 cipation of M. S. ENGEL was made possible by a Guggen- BECHLY, G., HAAS, F., SCHAWALLER, W., SCHMALFUSS,H.& heim Fellowship from the John Simon Guggenheim SCHMID, U. (2001): Ur-Geziefer – Die faszinierende Memorial Foundation (New York) and National Science Evolution der Insekten. – Stuttgarter Beiträge zur Natur- Foundation (USA) grant DEB-0542909. We are also grate- kunde, (C), 49: 96 pp. ful to HERBERT AXELROD for donating his large collection BEDFORD, G. O. (1978): Biology and ecology of the of Crato Formation fossils to the American Museum of Phasmatodea. – Annual Review of Entomology 23: 125- Natural History (New York). We thank HELMUT TISCH- 149. LINGER (Stammham, Germany) for numerous helpful com- BÉTHOUX,O.&NEL, A. (2002): Venation pattern and ments of an early draft of this manuscript and for providing revision of Orthoptera sensu nov. and sister groups. important literature, and GÜNTER VIOHL (Jura-Museum, Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu Eichstätt, Germany) and HELMUT MAY R (BSPGM, Mün- nov. – Zootaxa, 96: 1-88. chen, Germany) for give us the possibility to study the BERNER, R. A. (1998): The carbon cycle and CO2 over Chresmoda material housed in their museums. Thanks Phanerozoic time: the role of land plants. – Philoso- to DIMITRI E. SHCHERBAKOV (Laboratory of Palaeoento- phical Transactions of the Royal Society of London, (B), mology, Moscow, Russia) for give us the photograph of 353: 75-82. Saurophthirodes mongolicus. Research was carried out BEUTEL,R.G.&GORB, S. N. (2001): Ultrastructure of with the support of the Spanish Ministry of Science and attachment specializations of hexapods (Arthropoda): Technology CGL2005-00046/BTE. We thank SVEN evolutionary patterns inferred from revised ordinal BRADLER (University of Göttingen, Germany), DAV I D phylogeny. – Zeitschrift für Zoologische Systematik und GRIMALDI (AMNH, New York) and GÜNTER SCHWEIGERT Evolutionforschung, 39 (4): 177-207. 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WILLMANN, R. (2003): Die phylogenetischen Beziehungen Dr. A. NEL, CNRS UMR 5202, CP 50, Entomologie, der Insecta: Offene Fragen und Probleme. – Muséum National d’Histoire Naturelle, Paris, France; Verhandlungen Westdeutscher Entomologentag, 2001: e-mail : [email protected] 1-64. Dr. D. AZAR, Department of Biology, Faculty of Sciences II, ZOMPRO, O. (2001): The Phasmatodea and Raptophasma n. Lebanese Univ., Fanar-Matn, Lebanon; gen., Orthoptera incertae sedis, in Baltic amber (Insecta: e-mail: [email protected] Orthoptera). – Mitteilungen aus dem Geologisch- Dr. G. BECHLY, Staatliches Museum für Naturkunde Stutt- Paläontologischen Institut der Universität Hamburg, 85: gart, Rosenstein 1, D-70191 Stuttgart, Germany; 229-261. e-mail: [email protected] – (2004): Review of the genera of the Areolatae, including the status of Timema and Agathemera (Insecta, Phasma- Dr. J. A. DUNLOP, Museum für Naturkunde der Humboldt- todea). – Abhandlungen des Naturwissenschaftlichen Universität zu Berlin, D-10115, Berlin, Germany; Vereins in Hamburg, (N.F.), 37: 327 pp. e-mail: [email protected] – (2005): Inter- and intra-ordinal relationships of the Dr. M. S. ENGEL, Division of Invertebrate Zoology, Mantophasmatodea, with comments on the phylogeny of American Museum of Natural History, Central Park West polyneopteran orders (Insecta: Polyneoptera). – Mi- at 79th Street, New York, NY 10024-5192, and Division of tteilungen aus dem Geologisch-Paläontologischen Entomology (Paleoentomology), Natural History Museum, Institut der Universität Hamburg, 89: 85-116. and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Law- Manuscript received: September 21st, 2007. rence, KS 66049-2811, USA; Revised version accepted by the Stuttgart editor: December e-mail: [email protected] 14th, 2007. S. W. HEADS, Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Ports- Addresses of the authors: mouth, Burnaby Road, Portmsouth PO1 3QL, United Kingdom; Dr. X. DELCLÒS (to whom reprint requests should be addressed), Departament Estratigrafia, Paleontologia i Geo- e-mail: [email protected] ciències marines, Faculty of Geology, 08028 Barcelona, Spain; e-mail: [email protected]

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