Rock Crawlers in Baltic Amber (Notoptera: Mantophasmatodea)

2006·1.2 MB·English

by  Antonio Arillo

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3539, 10 pp., 5 figures, 2 tables December 7, 2006
Rock Crawlers in Baltic Amber
(Notoptera: Mantophasmatodea)
ANTONIO ARILLO1 AND MICHAEL S. ENGEL2
ABSTRACT
A fourth species of rock crawler (Notoptera: Mantophasmatodea: Mantophasmatidae) is
described and figured from an individual preserved in middle Eocene (Lutetian) Baltic amber.
AdicophasmagrylloblattoidesArilloandEngel,newspecies,isdistinguishedfromitscloserelative,
A. spinosum Engel and Grimaldi (reinstated), by the reduced pedicel, absence of spines on the
maxillae, absence of mesofemoral spination, and proportions of the thoracic segments. The
fossil shares with A. spinosum the presence of profemoral spination (confirmed by a new
photograph of the holotype) and absence of the dorsal profemoral carina, characters that
differentiate Adicophasma from the monotypic Raptophasma; it shares with all Baltic amber
Notoptera the absence of the setal fan on the arolium. As noted by previous authors, the
former order Mantophasmatodea is related to modern Grylloblattodea, whereas Mesozoic and
Paleozoic grylloblattodeans represent a stem group to both. As such, Grylloblattodea and
Mantophasmatodea are considered suborders of a single order, Notoptera Crampton (sensu
novum),followingthe recommendationofEngel andGrimaldi(2004).The namesforthree rock
crawlersareemendedinorderthatthespecificepithetmaymatchthegenderofthegenericname:
A. spinosum, Mantophasma zephyrum Zompro et al., and Tanzaniophasma subsolanum (Zompro
et al.) (nomina emendata). Raptophasmatinae and Ensiferophasmatidae are new synonyms of
Mantophasmatidae, while Tanzaniophasmatidae and Austrophasmatidae are newly demoted in
ranktoasubfamilyandtribeofMantophasmatidae,respectively.Ahierarchicalclassificationof
Polyneopterais appended.
1Departamento de Zoolog´ıa y Antropolog´ıa F´ısica (Entomolog´ıa), Facultad de Biolog´ıa, Universidad Complutense,
28040Madrid,Spain([email protected]).
2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;DivisionofEntomology(Paleoentomology),
Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive-Suite 140,
UniversityofKansas,Lawrence,Kansas66049-2811([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2006 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3539
INTRODUCTION Klass et al. (2002) were inconclusive in
regard to the affinities of the Mantophas-
The recent proposal of a new order of matodea, although they noted some similari-
moderninsects(Klassetal.,2002;Klass,2004) ties with the ice crawlers of the Northern
has stirred a great deal of attention in Hemisphere (order Grylloblattodea). Engel
entomology (e.g., Hall, 2002; Hansen, 2002; and Grimaldi (2004) further elaborated upon
Anonymous, 2003; Walker, 2003) as well as the affinity of Mantophasmatodea and
controversy (e.g., Tilgner, 2002; Klass, 2002). Grylloblattodea and even demonstrated the
The first mention of these insects appears to strong possibility that the two were sister
have been by Arillo et al. (1997), who groups. These same authors later further
providedthedescriptionofanenigmaticinsect indicated that Paleozoic and Mesozoic ‘‘gryl-
in Baltic amber, but did not name it. loblattodeans’’ represented stem groups to
Understandably, Arillo et al. left the fossil modern Mantophasmatodea + Grylloblat-
unassignedastoorder,discussingitsapparent todea and that features such as the loss of
affinities to Phasmatodea and other lineages. wings and ocelli were, in fact, additional
Zompro (2001) later described, albeit inade- synapomorphies of the two ‘‘orders’’
quately, additional material that was clearly (Grimaldi and Engel, 2005). Indeed, more
allied to Arillo et al.’s species, assigning the recent molecular studies have further sup-
taxon to the then new genus Raptophasma.3 ported the phylogenetic relationship between
Although Zompro (2001) formally considered Grylloblattodea and Mantophasmatodea4
Raptophasma as Orthoptera incertae sedis, he (Terry and Whiting, 2005). Following the
noted that the acquisition of modern species recommendation of Engel and Grimaldi
similartothefossilindicatedthepossibilityof (2004) we here combine the Mantophasma-
distinct ordinalassignment andevenprovided todeaandGrylloblattodeaintoasingleorder,
a name for the group as Raptophasmatodea employing the name Notoptera as proposed
(thus, technically the Mantophasmatodea was byCrampton(1915)forthecompositelineage.
notaneworderin2002asithadalreadybeen TheclassificationofNotopteraissummarized
named!). More recently, a third species was in table 1 (a classification of Polyneoptera is
established and placed in a new, putatively briefly outlined in appendix 2).
basal family (Zompro, 2005). Herein we provide a new and formal
Careful study of the modern material by description of the ‘‘first’’ rock crawler (i.e.,
Klaus-Dieter Klass revealed numerous addi-
tional characters, providing greater clarity 4EnigmaticallyTerryandWhiting(2005)statedthatthe
into the affinities of the insects than was molecular data supported the ordinal rank for
possible from the fossil material (Klass et al., Mantophasmatodea.Theapplicationofranksissemantic
andusedasareferencepointforrelativeinclusivenessof
2002). At that time two Recent species were
particular taxa. As such the data neither support nor
described and assigned to the genus Manto- refutetheapplicationofanyparticularrankforataxon.
phasma, and the order was formally proposed ThedatasimplyindicatethatmodernMantophasmatodea
(this time as Mantophasmatodea); the genus andGrylloblattodea(regardlessofrank)aresistertaxa.It
ismostconservativeandrational,giventhattheMesozoic
was described from species residing in
and Paleozoic fossils are basal to the combined rock
Namibia and Tanzania (Klass et al., 2002; crawler and ice crawler clade, to consider these taxa as
Zompro et al., 2002, 2004), but was shortly suborders of a single order encompassing the fossil and
thereafter recorded from the Karoo in South modern species (thereby avoiding the unnecessary pro-
liferation of orders to accommodate the stem-group
Africa (Picker et al., 2002). Engel and
species). These same authors superfluously provided
Grimaldi (2004) described a second fossil of anothernameforthisclade,Xenonomia,anunfortunate
the order from Baltic amber, Adicophasma name that refers to the malformation of the subordinal
spinosum (see appendix 1), and discussed the names Mantophasmatodea and Grylloblattodea rather
thananyanatomicaltraitunitingthegroups.Asnotedin
phylogeneticaffinitiesofthegroupasawhole.
the text, we follow the earlier proposals by Engel and
Grimaldi (2004) and Grimaldi and Engel (2005) of
3A re-examination and detailed descriptions of the expanding the far more apt name Notoptera for this
variousparatypesofR.kerneggerishouldbeundertaken order of insects and use the common name ‘‘crawler’’
toascertainwhethertheyhavebeencorrectlyassignedor (‘‘rock crawlers’’ for the suborder Mantophasmatodea
representotherspecies. and‘‘icecrawlers’’forthesuborderGrylloblattodea).
2006 ARILLO AND ENGEL: BALTICAMBER ROCKCRAWLERS 3
TABLE1
Hierarchical Classification of Notoptera
(Interfamilialclassification provided onlyfor Mantophasmatodea)
ORDERNOTOPTERACRAMPTON,1915
Family{TillyardembiidaeZalessky,1938
Family{BlattogryllidaeRasnitsyn,1976
SuborderGrylloblattodea1Brues&Melander,1932
FamilyGrylloblattidaeWalker,1914
SuborderMantophasmatodeaZomproetal.InKlassetal.,2002,stat.nov.
FamilyMantophasmatidae2Zomproetal.InKlassetal.,2002
Genus{RaptophasmaZompro,2001
Genus{AdicophasmaEngel&Grimaldi,2004
Genus{EnsiferophasmaZompro,2005
SubfamilyTanzaniophasmatinaeKlassetal.,2003,stat.nov.
GenusTanzaniophasmaKlassetal.,2003
SubfamilyMantophasmatinaeZomproetal.InKlassetal.,2002,stat.nov.
TribeTyrannophasmatiniZompro,2005
GenusPraedatophasmaZompro&AdisInZomproetal.,2002
GenusTyrannophasmaZomproInZomproetal.,2004
TribeMantophasmatiniZomproetal.InKlassetal.,2002,stat.nov.
GenusMantophasmaZomproetal.InKlassetal.,2002
GenusSclerophasmaKlassetal.,2003
TribeAustrophasmatiniKlassetal.,2003,stat.nov.
GenusAustrophasmaKlassetal.,2003
GenusHemilobophasmaKlassetal.,2003
GenusKaroophasmaKlassetal.,2003
GenusLobophasmaKlassetal.,2003
1BruesandMelander(1915)originallyconsideredthisgroupattheordinalrankbutemployedasuperfamilialsuffix(as
Grylloblattoidea).ThesesameauthorslateremendedthenametoGrylloblattodea,andwehaveusedthissubsequentdate
fortheoriginofthename.
2The usage of the family-group name Raptophasmatidae by Janzen (2002) is a nomen nudum (see also Engel and
Grimaldi,2004).ThenamewasvalidatedbyZompro(2005),butishereinnewlysynonymized,asitwasbasedsolelyon
plesiomorphiesandisparaphyletic(seeComments).
the specimen studied by Arillo et al., 1997: andthemoreelongateabdomen(seealsoKey
fig. 1). The age and origin of Baltic amber is to Baltic Amber Notoptera, below).
reviewed in Weitschat and Wichard (2002). DESCRIPTION: Female. Total body length
The format for the description and morpho- (excluding antennae) 6.0 mm (additional
logical terminology follows that employed by metrics provided in Arillo et al., 1997).
Engel and Grimaldi (2004). Coloration uniformly light brown. Head
transverse. Face with sparse, simple setae;
integument impunctate. Compound eyes
SYSTEMATIC PALEONTOLOGY large, extending entire length of head, encom-
passing almost entire lateral area of head;
Adicophasma grylloblattoides, new species
ommatidia small and numerous. Vertex
straight. Gena impunctate. Malar space
figures 1–4
exceedinglyshort.Antennalsocketapparently
DIAGNOSIS: The new species differs from just below facial midlength and near com-
its congener, A. spinosum Engel and Grimaldi pound eye; scape about equal in size to
(seeappendix1),bytheshortenedpedicelthat pedicel; pedicel about as long as wide; 16
isaboutaslongaswide,theabsenceofmeso- flagellomeres elongate and with sparse setae.
femoralspination,theabsenceofspinesonthe Maxillary palp five-segmented, basalmost
cardines,theproportionsofthethoracicnota, palpomere smallest, approximately equal in
4 AMERICAN MUSEUMNOVITATES NO. 3539
Fig. 1. Photomicrograph of holotype of Adicophasma grylloblattoides Arillo and Engel, new species
(MCNA10686).
size to second palpomere; third, fourth, and setae on outer surface (figs. 2–4); tarsi pen-
fifth maxillary palpomeres elongate; third tamerous, basal four tarsomeres with euplan-
palpomere longest; combined lengths of tulae; tarsomere IV weakly bilobed such that
fourth and fifth palpomeres slightly longer origin of tarsomere V slightly recessed, tar-
than length of third palpomere; distinct setae somereVelongateandslender,bearingsimple
scattered on apical palpomeres; stipes broad; pretarsal claws and large, broad arolium
cardo without spines. Coxae elongate; tro- (fig. 4); arolium without fans of elongate
chanters short; pro- and mesotibiae slightly setae. Middle and hind legs similar in con-
longer than respective femora; metatibia struction and with sparsely scattered, simple
longer than tibiae of preceding segments. setae; mesofemur not as swollen as profemur
Procoxa with three stout spines; profemur and lacking spination. Thoracic nota longer
noticeably swollen, without dorsal carinae, than wide and each of approximately equal
with two longitudinal rows of stout spines length and width, with exceedingly sparse,
ventrally (distinct ventral surface formed simple setae; integument impunctate; thorax
between rows of spines as in A. spinosum); about as broad as posterior width of head;
inner surface of protibia with similar stiff, pronotum only slightly longer than mesono-
elongate, simple spines as well as elongate tum. Abdomen elongate with sides relatively
Fig. 2. Illustration of holotype of Adicophasma grylloblattoides Arillo and Engel, new species
(MCNA10686).
2006 ARILLO AND ENGEL: BALTICAMBER ROCKCRAWLERS 5
Fig. 3. Photomicrograph of holotype foreleg of Adicophasma grylloblattoides Arillo and Engel, new
species(MCNA10686).
Fig.4. Illustrationofheadandpro-andmesothoracicsegmentsofAdicophasmagrylloblattoidesArillo
andEngel,new species(MCNA 10686).
6 AMERICAN MUSEUMNOVITATES NO. 3539
Fig. 5. Photomicrograph of holotype of Adicophasma spinosum Engel and Grimaldi showing detail of
forelegs, particularly the presence of profemoral and protibial spination (similar, albeit shorter and less
stout,spines are alsopresent onthe midlegs; seeEngeland Grimaldi, 2004).
parallel, segments short, terga and sterna Zompro (2005) considered the holotype of
broader than long; terga and sterna with A. spinosum as a nymph solely on its smaller
scattered, simple setae. Cerci short, unseg- body size. However, A. spinosum has fully
mented, gently tapering to apices, covered developed genitalia indicative of an adult and
with numerous erect to suberect elongate has numerous significant features distinguish-
setae. ing it from R. kerneggeri as noted in the
HOLOTYPE: Female, Baltic amber (middle original description (Engel and Grimaldi,
Eocene), MCNA 10686 (fig. 1), deposited in 2004) and in the following key. Perhaps most
the Museo de Ciencias Naturales de A´lava notablytheadultofRaptophasmaissupposed
(Vitoria, Spain). to lack femoral spines, while they are excep-
ETYMOLOGY: Thespecificepithetisarefer- tionally well developed in Adicophasma.
encetothesimilarityandphylogeneticaffinity Zompro’sstatementtothecontraryisentirely
of rock crawlers to the ice crawlers (suborder erroneous as the photograph in Engel and
Grylloblattodea). Grimaldi (2004) was not taken in the focal
COMMENTS: The new species, like all fossil plane of the legs, yet the out-of-focus spines
Notoptera, lack the setal fan on the arolium, on at least one leg can nonetheless be
a feature synapomorphic for the modern, discerned; similar, albeit slightly weaker,
African species of Mantophasmatodea. like spines are obvious on A. grylloblattoides
Raptophasma kerneggeri Zompro the new (fig. 3). That the spines are indeed present in
specieslacksmesofemoralspinationbutother- A.spinosumisevidencedbyanewphotograph
wise has all of the features of the genus oftheholotype taken inthefocal planeofthe
Adicophasma as characterized by Engel and legs (fig. 5), demonstrating that statements as
Grimaldi (2004). totheabsenceofthesestructuresaremeantto
2006 ARILLO AND ENGEL: BALTICAMBER ROCKCRAWLERS 7
mislead. Nymphal forms of rock crawlers can _ Femora without spination; profemur with
havespinesinthelatestofnymphalstages,but paired, dorsal, longitudinal carinae (Genus
only when such features are similarly present Raptophasma) . .. .. R.kerneggeri Zompro
in the adult (i.e., the latest nymphal stages 3. Mesofemur with distinct, ventral spination;
pedicel elongate, at least three times longer
acquire adult-like femoral spination). Thus,
than wide; cardo with two stout, apical
even if the holotype of A. spinosum were
spines; pronotum slightly longer than wide,
a nymph (which the terminal structures in-
meso- and metanota each wider than long;
dicate is not the case), it could scarcely be
lateral margins of abdomen convex (i.e.,
considered conspecific with R. kerneggeri, abdomen roughly ovoid); compound eyes
which is supposed to lack such features with large ommatidial facets (see Engel and
entirely in the adult stage [as evidenced by Grimaldi, 2004).. .. . .. .. .. .. .. .. .. .
the fact that the adult holotype male lacks .. . .. .. . A.spinosum Engeland Grimaldi
such characters, as do, apparently, the para- _ Mesofemur without ventral spination; pedicel
types; since spines in at least one paratype short, about as long as wide; cardo without
spines;allthoracicnotalongerthanwideand
were entirely overlooked (fig. 5), the series
each of approximately equal size; abdomen
should be re-examined]. The same can be said
elongate; compound eyes with smaller and
forthedorsalprofemoralcarinaepresentinR.
morenumerousommatidial facets.. .. .. .
kerneggeributlackinginAdicophasma,among
.. . .. .. .. .. .. . A. grylloblattoides, n.sp.
other generic features (see Engel and
Grimaldi, 2004). Most descriptions and fig-
DISCUSSION
ures of Baltic amber rock crawlers are in-
adequate (e.g., Zompro, 2001, 2005) as evi-
The new species described herein attests to
denced by the oversight of obvious features
the as yet unstudied diversity of polyneopter-
such as the presence/absence of femoral
an (particularly of the Orthopterida, sensu
spination among paratypes and should not
Grimaldi and Engel, 2005) insects in Tertiary
be relied upon. For more extensive descrip-
ambers,evenfromthosefaunaeasintensively
tions and figures refer to Engel and Grimaldi
studied as Baltic amber. In addition, the
(2004) and Grimaldi and Engel (2005). As the
amberNotopterafurtherhighlighttheimpor-
subfamily Raptophasmatinae and the family
tance of fossils for understanding the phylo-
Ensiferophasmatidae are based entirely on
genetic relationships, biogeographic patterns,
plesiomorphies, they are herein newly synon-
and other evolutionary phenomena of mod-
ymized with Mantophasmatidae (new synony-
ern taxa. For example, the Tertiary amber
mies). The Tyrannophasmatini is also suspi-
Notoptera indicate that the suborder
cious and thus is only tentatively retained
Mantophasmatodea is of bipolar distribution
(table 1). Indeed, the recognition of multiple
rather than a strictly Gondwanan group that
subfamilies or tribes within this remarkably
might be a relic of the rifting of the southern
small and relatively homogeneous family is
continents. Indeed, the presence of such
unwarranted.
amber fossils indicates that the suborder
Mantophasmatodea had a much wider distri-
bution as recently as the middle Eocene,
KEYTO BALTIC AMBER NOTOPTERA perhaps becoming extirpated from the
NorthernHemispherebythedramaticclimat-
1. Compound eyes occupying major portion of icshiftsoftheEocene-Oligocenetransition.It
head; antennae shorter than body; arolium is probable that Mantophasmatodea survived
projecting beyondpretarsalclaws . .. .. . 2 not only in southern Africa but in southern
_ Compound eyes placed in posterior half of
SouthAmericaandAsiaaswell.Ifthehabitat
head; antennae several times longer than
preferencesofthesouthernAfricanspeciesare
body; arolium not projecting beyond pre-
any indication, then suitable environmental
tarsal claws(GenusEnsiferophasma). .. ..
conditions may also be found in regions such
. .. .. .. .. . .. .. .E. velociraptor Zompro
2. At least some femora with distinct, ventral as Chile or Argentina. Indeed, given that
spination; profemur without dorsal carinae nymphs of rock crawlers may be generally
(Genus Adicophasma). .. .. .. .. . .. .. . 3 mistaken for immature Orthoptera or
8 AMERICAN MUSEUMNOVITATES NO. 3539
Phasmatodea, it is possible that the lineage Arillo, A., V.M. Ortun˜o, and A. Nel. 1997.
may eventually be discovered outside of Description of an enigmatic insect from Baltic
Africa. Similarly, rock crawlers may also be amber.BulletindelaSocie´te´Entomologiquede
France 102(1):11–14.
found in other fossiliferous ambers. It would
Brues, C.T., and A.L. Melander. 1915. Key to the
be particularly exciting to discover such taxa
families of North American insects: an in-
in Cretaceous ambers as these would un-
troduction to the classification of insects.
doubtedly provide powerful insights on the
Boston:Publishedbytheauthors,vii+140pp.
early history of Mantophasmatodea and
Brues, C.T., and A.L. Melander. 1932. Classifica-
perhaps also their sister Grylloblattodea as tion of insects: a key to the known families
they differentiated from stem-group families of insects and other terrestrial arthropods.
suchasBlattogryllidaeintheJurassicorEarly Bulletin of the Museum of Comparative
Cretaceous. Zoology73:1–672.
Crampton, G.C. 1915. The thoracic sclerites and
the systematic position of Grylloblatta campo-
ACKNOWLEDGMENTS
deiformis Walker, a remarkable annectant,
‘‘orthopteroid’’ insect. Entomological News
We are indebted to Dr. Torsten Wappler
26(10):337–350.
and an anonymous reviewer for careful com- Engel,M.S.,andD.A.Grimaldi.2004.Anewrock
mentsonanearlierversionofthemanuscript; crawlerinBalticamber,withcommentsonthe
to Dr. Bernd Hauser for correctly reminding order (Mantophasmatodea: Mantophasmati-
usofthegenderofPhasma;andtoDr. David dae).AmericanMuseumNovitates3431:1–11.
A. Grimaldi and Tam C. Nguyen for pro- Grimaldi, D., and M.S. Engel. 2005. Evolution of
viding the new, detailed photomicrograph of the insects. Cambridge: Cambridge University
Press, xv+755pp.
A. spinosum. This paper is an expansion of
Hall,R.2002.Nyinsektsordninghittad.Faunaoch
material presented at the 3rd International
Flora97(1):6–7.
Congress of Palaeoentomology (Pretoria,
Hansen, L.O. 2002. Mantophasmatodea, en ny
South Africa; February 2005). The participa-
insektorden funnet i Afrika. Insekt-Nytt 27(1–
tionoftheseniorauthor(AA)intheCongress
2):41–43.
was supportedby Diputacio´nForaldeA´lava, Janzen, J.-W. 2002. Arthropods in Baltic Amber.
whilebothauthorsaregratefultoDr.DenisJ. Halle:Ampyx Verlag,167pp.
Brothersfortheinvitationtoparticipateinthe Klass, K.-D. 2002. [Response to Tilgner, 2002].
meeting and hosting our visits to South Science 297:731a.
Africa. Thispaperisdedicatedtothememory Klass, K.-D. 2004. The new insect order Manto-
of Dr. James S. Ashe, whose tragic and phasmatodea(Insecta:Neoptera):morphology,
phylogenetic relationships and diversity.
untimely passing on 27 December 2005 has
Mitteilungen der Deutschen Gesellschaft fu¨r
left a hole in the hearts of not only the entire
Allgemeine und Angewandte Entomologie
UniversityofKansascommunitybutuponthe
14(1–6):63–66.
whole of entomology. This is contribution
Klass, K.-D., M.D. Picker, J. Damgaard, S. van
No. 3454 of the Division of Entomology, Noort, and K. Tojo. 2003. The taxonomy,
Natural History Museum, University of genitalic morphology, and phylogenetic rela-
Kansas. Partial support for the completion tionshipsofsouthernAfricanMantophasmato-
of this project was provided by NSF EF- dea (Insecta). Entomologische Abhandlungen
0341724 and DEB-0542909 (to MSE). 61(1):3–67.
Additional support was provided by (Accio´n Klass, K.-D., O. Zompro, N.P. Kristensen, and J.
Integrada con Francia HF2004-0053) and Adis. 2002. Mantophasmatodea: a new insect
order with extant members in the Afrotropics.
(Proyecto del Ministerio de Educacio´n y
Science 296(5572):1456–1459.
Ciencia de Espan˜a CGL2005-00046/BTE)
Picker,M.D.,J.F.Colville,andS.vanNoort.2002.
(both to AA).
Mantophasmatodea now in South Africa.
Science 297(5586):1475.
REFERENCES Rasnitsyn, A.P. 1976. Grylloblattids – present day
representatives of the order Protoblattodea
Anonymous.2003.It’sa…neworder!Rostrum60: (Insecta, Protoblattodea). Doklady Akademii
1–2. NaukSSSR 228(2):502–504.[In Russian]
2006 ARILLO AND ENGEL: BALTICAMBER ROCKCRAWLERS 9
Terry,M.D.,andM.F.Whiting.2005.Mantophas- gen aus dem Geologisch-Pala¨ontologischen
matodea and phylogeny of the lower neopter- Institut derUniversita¨tHamburg 85:229–261.
ousinsects. Cladistics 21(3):240–257. Zompro,O.2005.Inter-andintra-ordinalrelation-
Tilgner, E.H. 2002. Mantophasmatodea: a new ships of the Mantophasmatodea, with com-
insect order?Science 297:731a. ments on the phylogeny of polyneopteran
Walker, E.M. 1914. A new species of Orthoptera, orders (Insecta: Polyneoptera). Mitteilungen
forming a new genus and family. Canadian ausdemGeologisch-Pala¨ontologischenInstitut
Entomologist46: 93–99. der Universita¨tHamburg 89:85–116.
Walker, J.A. 2003. Mantophasmatodea – a new Zompro, O., J. Adis, P.E. Bragg, P. Naskrecki, K.
order of insects. Bulletin of the Amateur Meakin, M. Wittneben, and V. Saxe. . A new
Entomologist’s Society62(447): 72–78. genus and species of Mantophasmatidae
Weitschat, W., and W. Wichard. 2002. Atlas of (Insecta: Mantophasmatodea) from the
plants and animals in Baltic amber. Munich: Brandberg Massif, Namibia, with notes on
FriedrichPfeil, 256pp. behaviour. Cimbebasia 19: 13–24.
Zalessky, G.M. 1938. Nouveaux insectes permiens Zompro, O., J. Adis, and W. Weitschat. 2002. A
del’ordredesEmbiodea.AnnalesdelaSocie´te´ review of the order Mantophasmatodea (In-
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APPENDIX 1
A NOMENCLATURAL NOTE
As we were rightly reminded by our colleague Dr. Bernd Hauser (Muse´um d’histoire naturelle, Geneva),
Phasma is of neuter gender in Greek, and accordingly those names based upon this generic stem should
also be neuter (i.e., all of the currently recognized genus-group names in Mantophasmatodea). This
correctioningendernecessitatestheemendationofseveralspecificepithetsforrockcrawlersinordertobring
thespecificandgenericnamesintoalignment.Thefollowingemendationsarethusrequired(inalphabetical
order):
Adicophasmaspinosum Engeland Grimaldi, nomenemendatum
MantophasmazephyrumZomproet al.,nomenemendatum
Tanzaniophasmasubsolanum (Zompro etal.), nomenemendatum
10 AMERICAN MUSEUMNOVITATES NO. 3539
APPENDIX 2
HIERARCHICAL CLASSIFICATIONOF POLYNEOPTERA
Thefollowingtablebrieflysummarizesthehierarchicalclassificationofpolyneopteraninsects(detailswithin
the Dictyoptera are not outlined). Much work remains on the higher classification of this group and so
changes are to be expected, particularly with respect to the various fossil lineages. For example
Protorthoptera presently comprises taxa that are stem-group forms of various lineages across the
Polyneoptera(e.g.,manyarestem-groupOrthopterida,somestem-groupPlecopterida,&c.).Similarly,older
taxasuchasPermoplecoptera,Protelytroptera(inpart),andAeroplanopteraaregradestomodernlineages
suchas Plecoptera, Dermaptera, andPhasmatodea, respectively.
SupercohortPOLYNEOPTERAMartynov
‘‘{Protorthoptera’’Handlirsch,partim
CohortDictyopteraLeach
CohortAnartiopteraEngel
MagnorderPolyplecoptera,novum
SuperorderPlecopteridaBoudreaux
‘‘{Permoplecoptera’’Martynov
OrderPlecopteraBurmeister
MirorderMystropteraEngel
OrderZorapteraSilvestri
OrderEmbiodeaKusnezov
MagnorderPolyorthopteraEngel&Grimaldi
SuperorderDermapteridaBoudreaux
‘‘{Protelytroptera’’Tillyard,partim
OrderDermapteraDeGeer
SuperorderOrthopteridaBoudreaux
GrandorderNotopterodea,novum
OrderNotopteraCramptona
GrandorderPanorthopteraCrampton
+variousextinctstemgrouplineages
Order{GlosselytrodeaMartynov
Order{CaloneurodeaHandlirsch
Order{TitanopteraSharov
OrderOrthopteraOlivier
MirorderHolophasmatodeaGrimaldi&Engel
‘‘{Aeroplanoptera’’Tillyard
OrderPhasmatodeaBrunnervanWattenwyl
aThe clade consisting of the suborders Mantophasmatodea and Grylloblattodea, excluding the stem-group families
(e.g.,Tillyardembiidae),istermedNeonotopteraEngel,novum.