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A New Upper Cretaceous Species of Chresmoda from Lebanon

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A New Upper Cretaceous Species of Chresmoda from Lebanon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ig. 1. Chresmoda libanica sp. n., holotype specimen NI 3a, Fig. 2. Chresmoda libanica sp. n., holotype specimen NI 3a, photograph of general habitus. drawing of general habitus, scale bar represents 5 mm. 0.2 mm wide, bearing about 20 long and sharp spines long. The prothoracic femora are straight and elongate, (Fig. 10), probably corresponding to maxillary palpi. 10.7 mm long, 0.45 mm wide, bearing a short anterior These structures are better preserved in specimens NI 1 process and a series of small apical spines. The protho- and NI 2 than in NI 3. A short two- or three-segmented racic tibiae have a curved base, very short, 2.5 mm long, labial palpus is visible in NI 3, with no visible setae, 0.4 mm wide, representing the 23% of femoral length. about 1.0 mm long; mandibles visible in NI 3, 0.7 mm There is a row of small spines at tibial apex. The whole long and 0.5 mm wide, bearing a row of small obtuse prothoracic tarsus is 10.0 mm long, first tarsomere 3.0 teeth and several short but sharp spines (Figs 3–4). The mm long, second tarsomere about 2.5 mm long, third eyes are visible in NI 3, large elliptic, about 1.2 mm long about 1.0 mm long. The more distal segmentation of and 0.8 mm wide, 0.8 mm apart. The antennae are not visible, probably not preserved. The thorax is rather rounded but with a strong posterior concavity in ventral view, 4.0 mm long, 5.4 mm wide (NI 3), 2.5 mm long, 4.0 mm wide (NI 1), 2.5 mm long, 4.8 mm wide (NI 2), with no visible wing sheaths. The legs are very long and narrow, covered with very small hairs arranged in rows (visible in NI 2). The pairs of coxae are widely separated, 2.5 mm apart (NI 3), all similar, those of the same side of thorax being very close to each other, rounded and large, 0.8 mm wide and long. The trochanters are small and short, all similar, 0.5 mm Fig. 3. Chresmoda libanica sp. n., holotype specimen NI 3a, photograph of head and thorax, arrow indicates right mandible Fig. 4. Chresmoda libanica sp. n., holotype specimen NI 3a, with an apical short spine. drawing of ventral view of the body. Scale bar represents 5 mm. 146 . B1* >",,4 1 & !H' & RB 1 @ # & & !+ 5 5M> & &, HH % & IM4 ' ? I & , > & ' % &H>F & % &# !> , & + " & & . & , I & >,MH, & & ' & ' 9-!""!# & !+ , &, > % 9- !""!# & > > 7 & , 5H % & 5? ?F % = & % " > &>,MH, / ' ' %% & & % & & ' $ % = & & % & & % % &. & ' = % . U = %& C5#? + & &> > %C5#H , &5 ! %C ' ' & & % !# > , & + > % C +# % '0 =!">"E !#&& & = ' & , H C & , !4 % & 5+F & & % C 5 % N! + I & & '# & , 5 % & & % !4 & 1 = !"4?E N+ N!, ? & !!,M!!!.& 4> !,@& +# ' , 5 % N5 ' N!, + & , 5 %@&HM?# ' = % ' " ' % ' & 9 ' = % C &' ' & ' & 5?F ' ' & & & 9- !""!< '# & @= & !"">E @& +IH# 2 ' +,,+E+H?# 1 = !"4? L= = % & 0 ' 3 0 ' !""+# & # ' O / ' % & P 3 !45"# @ % . % ' = % & B !",?M!",4# 2 ' +,,+# '1 = !"4?E O U' & !,@& + # & !4I> V ' ' P & % / % & ' % 1 ' . % ' ' ' B U % = & .#'#!!'34!5!","#! ' +5F & >5F ' . # 9-!""!E@& ? ? 4# & .' ' C! C+ ' & 1 = !"4HE !>,# = % ' % & && ' ' 9- % & % ' !""!E>H+# $. * & & 16 Fig. 5. Chresmoda libanica sp. n., holotype specimen NI 3a, photograph of ovipositor. this last point. Interestingly, Martínez-Delclòs (1991) demonstrated that adult males of C. aquatica and C. obscura were apterous, suggesting that also the adult females of C. libanica could have been apterous. The presence of unique, highly specialized tarsi in C. obscura, C. aquatica, C. libanica sp. n., and probably C. orientalis strongly supports the hypothesis that these insects were living on the water surface, using the water superficial tension to “skate” on surface like extant Ger- Fig. 6. Chresmoda libanica sp. n., holotype specimen NI 3a, ridae and Veliidae, as already proposed by Martínez- drawing of ovipositor, scale bar represents 1 mm. Delclós (1991). Baudoin (1980) tried to establish that Chresmoda obscura could not live on water surface, (Baudoin 1980: 2). The oblique apices of the tarsomeres using the water superficial tension, through a comparison probably acted as a blocking system in order to avoid the with the extant Gerridae. He supposed that the Chres- complete distortion of the legs (see Figs 7–8, 13). Simi- moda were too large to float like Gerridae. But nobody larly, some Gerridae such as Potamobates thomasi Hun- knew in 1980 that at least some Chresmoda spp. could gerford, have no specialized setae but their have highly specialized legs and tarsi. Numerous extant two-segmented tarsi are curved (Andersen, 1982: Fig. Gerridae bear highly specialized setae that help them to 438). float by increasing their surface in contact with water sur- Nevertheless, the Chresmodidae probably did not use face (Andersen, 1982). Chresmoda aquatica and Chres- their legs for skating in the same way as modern Gerridae moda libanica sp. n. have no specialized setae on their and Veliidae do. These insects have short to very short legs but their exceptionally high number of tarsomeres fore legs, useless in skating, and they mainly use their (about 40 in C. libanica sp. n., about twelve in C. obscura median and partly hind legs for this purpose. The fore, and C. aquatica) probably helped to increase, thanks to mid and hind legs of the Chresmodidae have nearly the the curvature of the tarsi, the length of that part of the leg same lengths and structures, and so probably similar func- in contact with the water surface, and consequently to tions in skating. increase the vertical component of the superficial tension C. aquatica, Saurophthiroides mongolicus, and the Chresmodidae from Araripe (Lower Cretaceous of Brazil) probably lived in lacustrine palaeoenvironments. The Solnhofen lithographic limestone was a marine deposit, but it has yielded numerous fossils of terrestrial origin. The Nammoura “fish beds” are also of marine origin Fig. 8. Chresmoda libanica sp. n., holotype specimen NI 3a, Fig. 7. Chresmoda libanica sp. n., holotype specimen NI 3a, drawing of left median tibia and tarsi, scale bar represents 3 photograph of left median tarsus, arrows indicate tarsomeres. mm. 148 Fig. 10. Chresmoda libanica sp. n., paratype specimen NI 2b, photograph of maxillary palpi, arrow indicates some of the long and sharp spines of a palpus. less, two of the three known specimens of C. libanica sp. n. are well-preserved nymphal exuviae, which are very delicate remains that do not support long transport. This Fig. 9. Chresmoda libanica sp. n., paratype specimen NI 2b, suggests that these nymphs were living in the sea or very photograph of general habitus. close to it. C. libanica sp. n. was probably carnivorous since it (Schram et al., 1999), but with “a considerable influx of shows well-developed maxillary palpis, as in C. aquatica, terrestrial organic debris” (Krassilov and Bacchia, 2000: bearing numerous strong and sharp spines and it has 798). Thus both C. obscura and C. libanica sp. n. could strong mandibles. It would probably eat small insects have been marine or freshwater insects. trapped by the water surface tension, or planktonic or The long ovipositor with indentations
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