Research 63 (2016) 45e53

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Cretaceous Research

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A thorny, ‘anareolate’ stick- ( s.l.) in Upper Cretaceous amber from Myanmar, with remarks on diversification times among

* Michael S. Engel a, b, , Bo Wang c, d, Abdulaziz S. Alqarni e a Division of Entomology, Natural History Museum, 1501 Crestline Drive e Suite 140, University of Kansas, Lawrence, KS 66045-4415, USA b Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence, KS 66045, USA c State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China d Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Science, Beijing 100101, China e Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, P.O. Box 2460, Riyadh 11451, Saudi Arabia article info abstract

Article history: A new fossil stick-insect (Phasmatodea) is described and figured from a male preserved in Cretaceous Received 7 January 2016 amber from Myanmar. Echinosomiscus primoticus Engel and Wang, gen. et sp. nov., is a robust, somewhat- Received in revised form compressed stick-insect bearing abundant acanthae over the head and body, and remarkably lacks an 26 February 2016 area apicalis on the tibiae. The is described and assigned to a new, extinct of Phas- Accepted in revised form 27 February 2016 matidae s.l., as Echinosomiscinae Engel, subfam. nov. Brief remarks are made regarding the dating of Available online 3 March 2016 phasmatodean lineages, with E. primoticus providing the first reliable evidence for Euphasmatodea and even Neophasmatodea in the . Keywords: © Cenomanian 2016 Elsevier Ltd. All rights reserved. Euphasmatodea Holophasmatodea Neophasmatodea Phasmatidae

1. Introduction whereby gentle swaying gives the impression of foliage moving in a breeze (Bedford, 1978). The eggs of Phasmatodea are remarkably of the Phasmatodea are famous for crypsis, character-rich and mimic seeds, and the order is one of the few assuming diverse morphologies that allow them to mimic the with a developed ootaxonomy (Clark, 1976a, 1976b, 1978, 1979; surrounding foliage (Bedford, 1978), and their affinities among Sellick, 1988, 1997a, 1997b, 1998), making it possible to diagnose other polyneopteran lineages continues to inspire debate, either as the presence of particular from eggs alone. relatives to the Embiodea (Kristensen, 1975; Whiting et al., 2003; Naturally, in a group as varied as Phasmatodea, not all species Terry and Whiting, 2005; Friedemann et al., 2012) or Orthoptera can simply be lumped as either stick or leaf mimics, and there are (Hennig, 1981; Wheeler et al., 2001; Grimaldi and Engel, 2005). those, such as Trychopeplus Shelford or Oreophasma Günther that These prominent include some of the longest of extant are rather moss- or fern-like, and some Gray mimic insects, such as chani Bragg reaching to over 56 cm in lichens (Robinson, 1969; Brock, 2001). On the other end of the length when the legs are extended (Hennemann and Conle, 2008), spectrum, some forms are more robust and although long, are often and different groups are either greatly elongate, mimetic with compressed and not so clearly stick-like. Regionally known as land twigs or sticks, hence the name stick-insect, or are broadened and or tree ‘lobsters’, such phasmatodeans are often heavily armed by leaf-like, the aptly named leaf-insects of the (Phyllo- thorny projections, or acanthae (acanthae do occur widely phasmatodea). This extends also into their behavior, throughout Phasmatodea), and can be at times massive, reaching up to 15 cm and with a body masses nearing 25 g, although at masses up to 65 g females of the Malaysian Heteropteryx dilatata * Corresponding author. Division of Entomology, Natural History Museum, 1501 (Parkinson, 1798) tip the scales as the heaviest of Phasmatodea Crestline Drive e Suite 140, University of Kansas, Lawrence, KS 66045-4415, USA. E-mail addresses: [email protected] (M.S. Engel), [email protected] (Brock, 1999a). Perhaps the most famous of such insects is the (B. Wang), [email protected] (A.S. Alqarni). critically endangered (Montrouzier, 1855), the http://dx.doi.org/10.1016/j.cretres.2016.02.015 0195-6671/© 2016 Elsevier Ltd. All rights reserved. 46 M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53

Lord Howe Island Stick-Insect (sometimes referred to as the Lord long surface, beneath the insect, is comparatively flat, while the Howe Island Tree Lobster). Populations of this large, ‘Lazarus’ spe- opposing one is arched. The insect is oriented obliquely across the cies were decimated on Lord Howe Island by rats, becoming locally shorter axis (Fig. 1), with its face looking toward the curved rim of extinct by the 1960s, but in 2001 an isolated group of 24 individuals the oval. The legs are set alongside the body as if the were was rediscovered on Ball's Pyramid and it is considered the rarest of in stance, and the antennae are extended to the side and along the insects (Priddel et al., 2003; Robertson, 2006). Given their striking line of the body, although that of the left is incomplete. The morphologies and usually docile behavior, many stick- and leaf- is a bit shriveled, and the apicalmost segment partly insects are popular with zoological gardens and as pets for pri- damaged, the dorsomedial membrane torn, but preserving well vate collectors and children, and detailed guides exist to their the characteristic divided tenth tergum, albeit with the hemi- breeding and care (Brock, 1999a, 2000; Bradler and Seiler, 2012). tergites slightly twisted out of their natural position. The integu- The order is an ideal group from which to explore patterns of ment is partially cleared as preserved, and there is some Schimmel morphological evolution and convergence (e.g., Whiting et al., in places which, in combination with scattered organic debris 2003; Buckley et al., 2009), and is a textbook example of a group within the piece as well as the placement of the insect near some whereby wings are repeatedly lost and reacquired, presumably curved surfaces render various views difficult. Nonetheless, a through the turning on and off of a specific hox gene such that the remarkable amount of detail is observable, permitting a thorough genetic architecture for wings remains in place, albeit ‘dormant’, understanding of the species. even in those clades that are otherwise apterous. Parthenogenesis The mine from which the fossil was excavated is located in the abounds throughout Phasmatodea (Scali, 2009), and this repro- Hukawng Valley of Myanmar's northern state of Kachin, and near ductive mechanism appears to extend deeply in their phylogeny the village of Noije Bum, near Tainang, and westward of Myitkyina owing to its scattered phylogenetic presence across the order and (Kania et al., 2015). These mines have yielded well-studied occurrence in the relict Scudder thousands of fossils of earliest Cenomanian age (Shi et al., 2012), (Sandoval et al., 1998; Schwander et al., 2011), a group from west- and represent one of the richest and oldest sources for paleobio- ernmost North America isolated into its own suborder, Time- logical information on the Upper Cretaceous. The locality has been matodea, and sister to all other extant phasmatodeans, or extensively studied and mapped by Grimaldi et al. (2002) and Euphasmatodea (Bradler, 1999; Tilgner et al., 1999). Cruickshank and Ko (2003), and the biotic diversity, largely While the Phasmatodea include over 3000 modern species and comprising insects, has been tabulated by Grimaldi et al. (2002) are particularly diverse in southern and southeastern Asia (Brock, and Ross et al. (2010), but with numerous important additions 1999b; Seow-Choen, 2000; Bragg, 2001; Otte and Brock, 2005; subsequent to these works that have expanded the crustacean Chen and He, 2008; Großer,€ 2008), their past diversity has been a (Broly et al., 2015), arachnid (Engel and Grimaldi, 2014a; Engel challenge to reconstruct. Their often large proportions and frail et al., 2016a), and insect records (e.g., Grimaldi and Engel, 2013; bodies have not lent them to ideal preservation, and when fossils Barden and Grimaldi, 2014, 2016; Dikow and Grimaldi, 2014; have been recovered they are frequently of nymphs or exception- Engel and Grimaldi, 2014b; Grimaldi and Johnston, 2014; Arillo ally fragmented. Varied Mesozoic fossils from the and et al., 2015; Parker and Grimaldi, 2014; Vea and Grimaldi, 2015; Lower Cretaceous, many known only from isolated wings, have Delclos et al., 2016; Engel et al., 2016b, 2016c, 2016d, 2016e, been attributed to the stem group of Phasmatodea, the most 2016f; Yamamoto, 2016). notable being those of the Aeroplanidae, Aerophasmatidae, and The classification of Phasmatodea is rather muddled, with Susumaniidae (e.g., Tillyard, 1918; Sharov, 1968; Gorochov, 1993, much comparative morphological investigation and revision 1994; Ren, 1997; Gorochov and Rasnitsyn, 2002; Shang et al., necessary before a robust, stable system is achieved. The Bradley 2011; Wang and Ren, 2013; Wang et al., 2014 d the Chresmodi- and Galil (1977) system has repeatedly been found wanting (e.g., dae are no longer attributed to Holophasmatodea: Delclos et al., Bradler, 2001, 2009; Tilgner, 2002; Whiting et al., 2003; Buckley 2008). Collectively, these varied Mesozoic families constitute with et al., 2009), and while Günther's (1953) classification is older crown-group Phasmatodea a superordinal group known as the and certainly out of date, it is broadly more accurate. Phyloge- Holophasmatodea (Grimaldi and Engel, 2005), although some au- netic studies have revealed considerable paraphyly and polyphyly thors have doubted their relatedness to Phasmatodea (e.g., Ragge, (Tilgner, 2002; Whiting et al., 2003; Bradler, 2009; Buckley et al., 1955). While stick-like species are known from Cenozoic deposits 2009; Komoto^ et al., 2011; Bradler et al., 2014), although some- (Tilgner, 2000), and those documented Mesozoic forms are rather times in contradictory ways, reflecting a general lack of resolu- elongate, the more extreme morphologies of the phylliids lack a tion for even the most fundamental of divisions among fossil record outside of a single, remarkable species from the mid- Euphasmatodea. The older divisions of Areolatae and Anareolatae of Germany (Wedmann et al., 2007). (Stål, 1875; Brunner von Wattenwyl and Redtenbacher, 1906, Here we document a small, but robust, thorny stick-insect in 1907, 1908), are not natural (Bradler, 2001, 2009; Tilgner, 2002), amber from the Upper Cretaceous of northern Myanmar (Fig. 1). and therefore of no practical use, except to note that the presence This species is remarkable in that it clearly possesses the special- of the areola apicalis is putatively plesiomorphic for Phasmatodea ized acanthae so familiar in many lineages of Phasmatodea, (Bradler, 2009). We concur with Hennemann et al. (2009) and including a raised cephalic crown of spines. We provide a Conle et al. (2011), that the systems, largely of the ‘areolates’ but description of the approximately 98.8 Ma (Shi et al., 2012) fossil as also of other groups, of Zompro and Großer€ (2003) and Zompro an aid to understanding the history of Phasmatodea and the early (2004a, 2004b) are confused and do more to crimp work than appearance of these specialized morphologies. advance it. Accordingly, we generally follow the various re- organizations of the order as advocated by Hennemann and 2. Material and methods Conle (2008), Bradler (2009),andBradler et al. (2014). For the descriptions, we adopt generally elements of morphological ter- The thorny stick-insect was identified in a small, oblong, ovoid minology from Rehn and Rehn (1938 (1939)), Günther (1953), piece of light yellow-orange amber, and of about 18.5 mm in Beier (1968), Bragg (1997),andBradler (2003), and we present maximum length and 12 mm in maximum breadth. The piece is these accounts in the philosophy that solid descriptive science rather low, with a maximum depth near its center of about represents the foundation upon which evolutionary patterns and 4.5 mm, and tapering gently and radially toward its edges. One explanations are built (Grimaldi and Engel, 2007). Photographs M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53 47

Fig. 1. Microphotographs of holotype male (NIGP 163536) of Echinosomiscus primoticus Engel and Wang, gen. et sp. nov., in Burmese Cenomanian amber. A, Dorsal view as pre- served, viewed through curved upper surface of amber piece (scale bar ¼ 1 mm). B, Dorsolateral oblique view of cephalic acanthae (scale bar ¼ 0.25 mm). C, Dorsolateral oblique view of head and anterior portion of thorax (scale bar ¼ 0.5 mm).

were taken through an Infinity K-2 long-distance microscope lens (e.g., Hennemann and Conle, 2008). Although the general mounted to a Canon 7D digital camera, while measurements habitus of the present fossil is similar to some (Heter- were done with the aid of an ocular micrometer on an Olympus opterygidae) or some , the distinct absence of the SZX-12 stereomicroscope. All taxonomic actions established area apicalis on the tibiae (there are particularly clear views of this herein have been registered in ZooBank LSID urn:lsid:zoo- on the mesotibiae, but also on one of the metatibiae), clearly in- bank.org:pub:E279FC5E-CD94-44E0-B063-DBEF0115CEA6. dicates a placement outside of the areolate lineages. The combi- nation of the absence of areole on all tibiae, the comparatively long antennae, the absence of wings, and the division of the tenth 3. Systematic palaeontology abdominal tergum into moveable hemitergites are indicative of a placement among phasmatids, in the broad sense, and near the Family Phasmatidae Gray, 1835, s.l. traditional subfamily (sensu Günther, 1953) or the Comments. The definition of Phasmatidae is far from stable, with ‘Schizodecema’ (sensu Bradler, 2009: although this grouping may some authors preferring a considerably narrowed concept for the not monophyletic, e.g., Bradler et al., 2014). Certainly, the moveable 48 M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53 hemitergites with an apical, circular field of small, sclerotized tu- Etymology. The new genus-group name is a combination of the bercles (‘Dornenfeld’) is similar to that of the Lonchodinae and Greek, echinos, meaning, “spiny”; soma, meaning, “body”; and the Clitumninae (sensu Hennemann and Conle, 2008). However, there suffix iskos, generally denoting a diminutive. The gender of the is little resemblance between the present fossil group and these name is masculine. The generic name is registered under ZooBank more stick-like , as circumscribed, although with the LSID urn:lsid:zoobank.org:act:1D41C94E-7F1A-4725-A5DB- incorporation of various eurycanthines (sensu Bradler et al., 2014), 572D78A49EF8. lonchodines are a much more heterogeneous lineage and includes Echinosomiscus primoticus Engel and Wang, sp. nov. various robust genera with fewer antennomeres and more dentate (Figs. 1e4) legs. Nonetheless, the combination of characters embodied in the present fossil does not accord with Lonchodinae, even when Etymology. The specific epithet is taken from the Latin, primoticus, incorporating Eurycanthinae, and it is accordingly placed within its meaning, “happening first”, and refers to this being the earliest own subfamily pending a comprehensive revision of familial amber-preserved thorny stick-insect. The epithet is registered under groups within Neophasmatodea. ZooBank LSID urn:lsid:zoobank.org:act:7456107E-71C6-4E2A- 884A-318F6800069B. Echinosomiscinae Engel, subfam. nov. Holotype. _ (Fig. 1A), NIGP 163536; amber; lowermost Cenomanian Type genus: Echinosomiscus Engel, gen. nov. (Shi et al., 2012), Upper Cretaceous; mine near village of Noije Bum, Diagnosis. Small, apterous insects (approximately 7.9 mm in near Tanaing, Hukawng Valley, Kachin State, Myanmar. The holo- length), with a generally spinose body (Figs. 1A, 2A). Head sub- type is located in the collections of the Nanjing Institute of Geology globular (Figs. 1, 3A, 3B), only slightly prognathous, apparently and Palaeontology, Chinese Academy of Sciences, Nanjing, China. slightly wider than long, with vertex slightly raised along line of Diagnosis. As for the genus (vide supra). coronal acanthae (Figs. 1B, 1C, 3A, 3B); ocelli absent; compound Description. _: Total body length (as preserved) 7.9 mm; body and eyes hemispherical, exophthalmic; antenna filiform, longer than legs generally spinose, integument generally brown (except forelegs (Fig. 1A), extending in repose to at least apex of first portions of flagellomeres IeVI white) and smooth between abdominal tergum (‘median segment’); scape cylindrical, much spines. longer than wide, with two dorsal spines; pedicel cylindrical, short, less than one-third length of scape; eight flagellomeres, Head subglobular, apparently slightly wider than long, width with individual flagellomeres narrow elongate, apical flag- across compound eyes 1.76 mm (directly lateral or frontal view ellomeres progressively shorter, with apical two flagellomeres impossible as preserved and so a precise measure of head length much shorter than preceding flagellomeres. Thorax spinose could not be achieved); vertex slightly elevated above compound (Figs. 1A, 2A, 2B), with each segment wider than long; pronotum eyes and along line of coronal acanthae; vertex posterior to coronal shorter than mesonotum; mesonotum subequal to metanotum; acanthae short and sloping gently to rounded occiput; genae abdominal tergum I (‘median segment’) fused to metanotum, rounded, slightly longer than compound eyes; ocelli absent; broadly transverse, much shorter than metanotum; nota with compound eyes hemispherical, exophthalmic, with abundant prominent rows of anterior, posterior, and lateral spines as well as minute ocular setae; antenna filiform, longer than forelegs, spines on discs; pleura spinose; thoracic sterna broad, flattened; extending in repose to at least apex of abdominal tergum I (‘me- prosternum without specialized sensory patches; abdominal dian segment’); scape cylindrical, much longer than wide, length sternum I indistinguishably fused to metasternum. Legs spinose 0.67 mm, with two dorsal spines, first near midlength, second at (Figs. 1A, 2A, 2C, 3D), with carinae dentate, albeit femoral dorsal about apical quarter; pedicel cylindrical, short, length 0.17 mm; carinae somewhat weakened giving dorsal surfaces a somewhat eight elongate flagellomeres, each rounded in cross-section, nar- curved appearance; femora rectangular to weakly trapezoidal in row, and distinctly thinner than scape and slightly thinner than cross-section; profemur shorter than head and pronotum com- pedicel; flagellomeres of subequal widths; flagellomere I length bined, slightly curved basally, with anterior-ventral carina weakly 0.55 mm, width 0.08 mm; flagellomere II length 0.70 mm; flag- lamellate apically; metafemur thickened, heavily spinose, with ellomere III length 0.78 mm; flagellomere IV length 0.83 mm; anterior-ventral carina strongly lamellate in apical third, medi- flagellomere V length 0.75 mm; flagellomere VI 0.72 mm; flag- oventral carina absent; all tibiae lacking area apicalis; metatibia ellomere VII 0.41 mm; flagellomere VIII 0.48 mm; pedicel and slightly shorter than metafemur; tarsi (pentamerous) with dis- flagellum bearing numerous, long, fine, suberect setae; mandibles titarsomeres (tarsomere V) subequal to combined lengths of broad, massive; maxillary palpus elongate, pentamerous (five remaining tarsomeres; basitarsi slightly longer than tarsomeres II; palpomeres), projecting in front of head as preserved; labial palpus pretarsal claws large, simple. Abdomen short, narrow, with rows of short, trimerous (three palpomeres). Acanthotaxy of head capsule spines apically and laterally on terga, without lateral lobes; anal as follows: pair of small, inner orbital spines present near upper segment not tectiform; tenth tergum deeply divided into move- inner border of compound eye (Fig. 3A); anterior occipital acan- able, elongate hemitergites (Fig. 4), apically bearing on inner sur- thae large, with small paramedial spines along transverse frontal face a produced, circular patch of sclerotized tubercles, basally ridge between anterior occipital acanthae (Fig. 3A, 3B); coronal hemitergites apparently connected by thin sclerotized bridge and acanthae raised on an elevated, almost-lamellate ridge (Fig. 1B, 1C), broad membrane; vomer absent; cercus (consisting of single cer- paired lateral coronal acanthae larger than paramedial coronal comere), long, fingerlike (not compressed, not lanceolate or foli- acanthae (Figs. 1B, 1C, 3A, 3B). aceous) (Fig. 4). Thorax robust, broad, spinose (Figs. 2B, 3C), somewhat dorso- Remark. The subfamilial name is registered under ZooBank LSID ventrally compressed. Pronotum slightly longer and wider than urn:lsid:zoobank.org:act:4C5C075A-FFC6-43AC-B9AF- head, wider than long, medial length 0.80 mm, width 2.15 mm, 798CA5C6F7A5. roughly trapezoidal, widening abruptly to about midline, then widening more gradually to posterior border, anterior border rim- Echinosomiscus Engel, gen. nov. med and slightly raised, with weak transverse depression imme- Type species: Echinosomiscus primoticus Engel and Wang, sp. nov. diately posterior to anterior rimmed region, not reaching to lateral borders; surface without mediolongitudinal or lateral-longitudinal Diagnosis. As for the subfamily (vide supra). lines or carinae. Prosternum broad, flattened, widely separating M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53 49

Fig. 2. Microphotographs of holotype male (NIGP 163536) of Echinosomiscus primoticus Engel and Wang, gen. et sp. nov. A, Ventral view as preserved (scale bar ¼ 1 mm). B, Mesonotum, metanotum, and anterior part of first abdominal tergum (scale bar ¼ 0.5 mm). C, Left hind leg (scale bar ¼ 0.5 mm).

procoxae, apparently without specialized sensory patches. Acan- Metanotum about as long as mesonotum, wider than long, medial thotaxy of pronotum: rimmed anterior border of pronotum with length 1.05 mm, width 2.40 mm, lateral margin widening slightly, row of short spines; pronotal surface with median pronotal acan- then weakly tapering toward posterior. Metasternum broad, flat- thus; posterior border of pronotum with row of short spines; large, tened, widely separating metacoxae. Acanthotaxy of metanotum: paired, posterolateral acanthae present. Mesonotum slightly longer metanotum with anterior row of spines and prominent antero- than pronotum, wider than long, medial length 1.10 mm, width lateral acanthae; paired mediolateral acanthae and smaller 2.40 mm, lateral margin convex, widest at about midlength; surface posterior-lateral spine; with posterior row of spines. (Thoracic without longitudinal impressions or carinae. Mesosternum broad, pleura apparently with scattered spines but owing to nature of flattened, widely separating mesocoxae. Acanthotaxy of meso- preservation their precise number and location is impossible to notum: mesonotum with anterior row of spines, anterior marginal ascertain). Abdominal tergum I (‘median segment’) fused to meta- acanthus larger than spines of anterior row; anterolateral acanthus notum but with distinct line of fusion, transversely rectangular, prominent; mesonotal disc with medial row of spines; posterolat- about as wide as metanotum, much shorter than metanotum, eral mesonotal acanthus prominent; lateral mesonotal acanthae medial length 0.55 mm, posterior width 2.20 mm, with posterior prominent, anterior lateral paired, posterior lateral single. margin straight; with prominent anterolateral acanthus and strong 50 M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53

Fig. 3. Line drawings of holotype male (NIGP 163536) of Echinosomiscus primoticus Engel and Wang, gen. et sp. nov., in Cenomanian amber from northern Myanmar. A, Head in dorsal view (scale bar ¼ 1 mm). B, Head in oblique, upper frontolateral view. C, Thoracic nota and fused first abdominal tergum (‘median segment’) (scale bar ¼ 0.5 mm). D, Hind leg in lateral view (scale bar ¼ 0.5 mm). row of posterior spines (Fig. 3C); abdominal sternum I indistinctly fused with metasternum. Legs somewhat stout, strongly spinose over all podites (Figs. 1A, 2A, 2C, 3D) except tarsi and pretarsi; dorsal carinae weak on mid and hind legs, more distinct on forelegs; all femora rectangular- trapezoid in cross-section; all tibiae without area apicalis; profe- mur length 1.46 mm, protibia length 1.31 mm, protarsus length 1.22 mm; mesofemur length 1.50 mm, mesotibia length 1.25 mm, mesotarsus length 1.22 mm; metafemur length 2.33 mm, metatibia length 2.12 mm, metatarsus length 1.24 mm. Abdominal segments IIeIX distinctly narrower than thorax, abdomen slightly tapering in width apically from segments IIeVIII, with terga IIeVIII wider than long, apical margins straight; segment IX larger than preceding segments, with tergum IX slightly wider Fig. 4. Microphotographs of terminalia of Echinosomiscus primoticus Engel and Wang, than preceding tergum, nearly as long as wide, dorsal surface gen. et sp. nov. (scale bar ¼ 0.25 mm). A, Dorsal view. B, Ventral view. somewhat more convex, medioapical margin broadly concave, with prominent lateral and apical spines; tergum X elongate, divided into moveable hemitergites (the hemitergites are torsion-rotated 4. Discussion out of their natural position as preserved, indicative of their ease of movement; the right hemitergite is almost completely twisted There are many challenges in the placement of any fossil around and with considerable damage to its base; the left hemi- phasmatodean, their often specialized morphology belies phylo- tergite is only partially rotated inward: Fig. 4), apically with genetic affinities and as a result the current classification is vexa- rounded area bearing rounded field of sclerotized tubercles (thorn tious. As mentioned earlier, the extant families and subfamilies are field) on inner surface; hemitergites apparently mediobasally burdened with much paraphyly and polyphyly (e.g., Bradler, 2009; united by thin sclerotized bridge and membranous area (partially Buckley et al., 2009; Bradler et al., 2014). Without clearly defined shredded as preserved); cercus undivided, consisting of single clades and morphological synapomorphies for these groups, the cercomere, long, finger-like, tapering in width to acutely rounded placement of any fossil taxon is fraught with obstacles, particularly apex and bearing numerous elongate, stiff setae, length 0.49 mm; true for incompletely preserved material, and this has left open abdominal sterna comparatively flat and without spines. many questions regarding the interpretation of numerous putative \: Latet. records (Tilgner, 2000). At the least, it is abundantly evident that M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53 51

Phasmatodea are monophyletic, and that the varied Mesozoic lin- estimation is exceedingly sensitive to calibration methods (e.g., eages hitherto reported form a grade of families that, along with Ware et al., 2010; Sauquet et al., 2012), and admittedly the fossil crown-group phasmatodeans, form the Holophasmatodea record of Phasmatodea as it has hitherto been documented leaves (Willmann, 2003; Grimaldi and Engel, 2005). The suborders few reliable records from which to calibrate any analysis. In this and Euphasmatodea (inclusive of the southern South regard, the young ages are largely influenced by the comparatively American Agathemera Stål), are reciprocally monophyletic and basal divergence of Phylliidae and the Lutetian age for Eophyllium sister groups (Sandoval et al., 1998; Bradler, 1999, 2009; Tilgner Wedmann et al. (2007), more so than any enriched documentation et al., 1999; Tilgner, 2002; Whiting et al., 2003). The putative of numerous phasmatodean lineages from the Cenozoic, and rests infraorder Agathemerodea is apparently nested among or near on the unfounded assumption that this fossil appeared immedi- Pseudophasmatinae (Günther, 1953; Whiting et al., 2003; Bradler ately at the divergence of the phylliid lineage. In fact, although et al., 2014), and should be abandoned, and concomitantly also crown-group phylliids (inclusive of Eophyllium), may be of early the unnatural infraorder Verophasmatodea (i.e., all Euphasmatodea Eocene age, this group could have an extended stem and diverged exclusive of Agathemera). The Aschiphasmatidae are putatively the from other Phasmatodea at a much earlier date. The genus basal branch of Euphasmatodea (Tilgner, 2002; Buckley et al., Eophyllium documents less the divergence of Phylliidae from other 2009), and comprise infraorder Aschiphasmatodea, while the Phasmatodea than it does simply the earliest appearance of the remainder of the order form the Neophasmatodea (i.e., all node comprising Eophyllium and other phylliids, which cannot be Euphasmatodea excluding aschiphasmatoids). The leaf-insects assumed to be equivalent in age to their extant sister . While (Phylliidae) are repeatedly recovered as natural (e.g., Buckley sister lineages are of the same age owing to their point of diver- et al., 2009; Bradler et al., 2014), and at least the families Aschi- gence, their corresponding extant crown groups may be of radically phasmatidae, s.str., and s.str. different ages depending on when the first subordinate clade appear to be monophyletic (Tilgner, 2002; Bradler et al., 2014). within each takes on recognizable apomorphies for the clade as we There are no fossil records for Timematodea, and although the circumscribe them. If there is an extended stem between the Baltic amber Electrotimema Zompro has been assigned to this ‘Phylliidae-other Phasmatodea’ divergence and the appearance of suborder, there is no evidence at present to support such a place- the ‘Eophyllium þ other Phylliidae’ node, then Eophyllium are not ment. Indeed, the two accounts of the sole species in this Eocene indicative of a calibration for the phylliid divergence, and corre- genus are not easily compared (measurements and descriptive spondingly, the first appearance of a crown-group branch for the details do not match between the two descriptions despite being ‘other Phasmatodea’ clade could have taken place much earlier prepared from the same specimen, putatively No.2632 of the than the origin of the ‘Eophyllium-other Phylliidae’ split. In order to former Grohn€ collection, now in the University Museum of accurately time the divergences of the various major events among Hamburg); for example, the earlier treatment figured and Phasmatodea it is clear that we require far more paleontological described five tarsomeres (Zompro, 2001), the latter mentioned material that can be adequately characterized and incorporated only the first three tarsomeres but never actually stated that the into phylogenetic work alongside modern genera, and ideally tarsi are trimerous (as is the case for Timematodea) (Zompro, incorporating those Mesozoic members of the Holophasmatodea 2005). The majority of fossil records for Euphasmatodea come (much as was attempted by Willmann, 2003). Regardless, Echino- from amber of Eocene to Miocene age (e.g., Tilgner, 2000), as well as somiscus primoticus provides an isolated but nonetheless early compressions from the Eocene of Europe (Lutz, 1990; Wedmann glimpse into the Cenomanian fauna of Phasmatodea and reveals a et al., 2007) and the remarkable possibility of a fossil Agathemera much earlier occurrence of Neophasmatodea. from the Florissant shales of Colorado (Scudder, 1890). An Eocene fossil from Oise amber attributed to Euphasmatodea (Nel et al., 5. Concluding remarks 2010), does not appear to belong to the order (Bradler and Buckley, 2011) and this particular fossil, which is would be of The discovery of Echinosomiscus primoticus, while not revealing particular interest to study via m-CT scanning in order to better of broad-scale patterns in the evolution of Phasmatodea, none- reveal its characters as it could be important for identifying a theless, demonstrates the early occurrence of the characteristic possible Phasmatodea-Orthopterida affinity. Fossils of phasmato- robust, somewhat flattened, and spiny morphology so familiar in dean eggs also document the occurrence of Euphasmatodea from many of the Southeast Asian fauna today. While such insects are various Cenozoic localities (Sellick, 1994; Poinar, 2011), and, quite often today large in size, their Cretaceous cousins were diminutive unexpectedly, in Burmese amber (Rasnitsyn and Ross, 2000). Un- and it is an open question as to what factors led to the gradual fortunately, this scarcity of material has rendered the fossil record increase in body size. The present fossil provides an important re- of Phasmatodea has less than illuminating, particularly in regard to cord for ascertaining the timing of events in stick-insect evolution the timing of divergence events and the appearance of particular and documents comparatively derived Euphasmatodea and even phasmatodean traits. Echinosomiscus primoticus assumes some Neophasmatodea in the earliest Cenomanian, although the pres- significance for its corroboration of stick-insects in the Cenomanian ently confounded state of phasmatodean classification and phy- fauna of northern Myanmar, and more importantly the presence of logeny is certainly a hindrance to much further resolution. putatively derived Euphasmatodea. In their molecular analysis Buckley et al. (2009), using as cali- bration points the Eocene phylliid (Wedmann et al., 2007) and the Acknowledgments report of Cenomanian phasmatodean eggs (Rasnitsyn and Ross, 2000), obtained a minimum age of 51.9 Ma for the euphasmato- We extend our thanks to the National Basic Research Program of dean radiation and considered most lineages therein as having China (2012CB821900), the National Natural Science Foundation of diverged between the Ypresian and Burdigalian (Miocene). If this China (41572010, J1210006), and the Youth Innovation Promotion were the case, then the Phasmatodea would represent one of the Association of the Chinese Academy of Science (No. 2011224) for youngest of insect orders to diversify, rivaled in its ‘infancy’ by only financial support of this work. We are further grateful to Ms. Jen- the true lice (Phthiraptera) (Grimaldi and Engel, 2005). The dis- nifer C. Thomas for expert assistance with microphotography, to covery of a phasmatid in the Cenomanian amber of Myanmar casts Mrs. Kellie K. Magill-Engel for encouragement throughout the doubt on these age estimations. Indeed, molecular-only divergence project, and to two anonymous reviewers for their helpful input. 52 M.S. Engel et al. / Cretaceous Research 63 (2016) 45e53

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