Adicophasma grylloblattoides, ARILLO & ENGEL, 2006

ARILLO, ANTONIO & ENGEL, MICHAEL S, 2006, Rock Crawlers in Baltic Amber (Notoptera: Mantophasmatodea), American Museum Novitates 3539 (1), pp. 1-10 : 3-7

Adicophasma grylloblattoides , new species figures 1–4

DIAGNOSIS: The new species differs from its congener, A. spinosum Engel and Grimaldi (see appendix 1), by the shortened pedicel that is about as long as wide, the absence of mesofemoral spination, the absence of spines on the cardines, the proportions of the thoracic nota, and the more elongate abdomen (see also Key to Baltic Amber Notoptera , below).

DESCRIPTION: Female. Total body length (excluding antennae) 6.0 mm (additional metrics provided in Arillo et al., 1997). Coloration uniformly light brown. Head transverse. Face with sparse, simple setae; integument impunctate. Compound eyes large, extending entire length of head, encompassing almost entire lateral area of head; ommatidia small and numerous. Vertex straight. Gena impunctate. Malar space exceedingly short. Antennal socket apparently just below facial midlength and near compound eye; scape about equal in size to pedicel; pedicel about as long as wide; 16 flagellomeres elongate and with sparse setae. Maxillary palp five-segmented, basalmost palpomere smallest, approximately equal in

(MCNA 10686).

size to second palpomere; third, fourth, and fifth maxillary palpomeres elongate; third palpomere longest; combined lengths of fourth and fifth palpomeres slightly longer than length of third palpomere; distinct setae scattered on apical palpomeres; stipes broad; cardo without spines. Coxae elongate; trochanters short; pro- and mesotibiae slightly longer than respective femora; metatibia longer than tibiae of preceding segments. Procoxa with three stout spines; profemur noticeably swollen, without dorsal carinae, with two longitudinal rows of stout spines ventrally (distinct ventral surface formed between rows of spines as in A. spinosum); inner surface of protibia with similar stiff, elongate, simple spines as well as elongate setae on outer surface ( figs. 2–4 ); tarsi pentamerous, basal four tarsomeres with euplantulae; tarsomere IV weakly bilobed such that origin of tarsomere V slightly recessed, tarsomere V elongate and slender, bearing simple pretarsal claws and large, broad arolium ( fig. 4 ); arolium without fans of elongate setae. Middle and hind legs similar in construction and with sparsely scattered, simple setae; mesofemur not as swollen as profemur and lacking spination. Thoracic nota longer than wide and each of approximately equal length and width, with exceedingly sparse, simple setae; integument impunctate; thorax about as broad as posterior width of head; pronotum only slightly longer than mesonotum. Abdomen elongate with sides relatively

(MCNA 10686).

parallel, segments short, terga and sterna broader than long; terga and sterna with scattered, simple setae. Cerci short, unsegmented, gently tapering to apices, covered with numerous erect to suberect elongate setae.

HOLOTYPE: Female, Baltic amber (middle Eocene), MCNA 10686 ( fig. 1 ), deposited in the Museo de Ciencias Naturales de Álava (Vitoria, Spain).

ETYMOLOGY: The specific epithet is a reference to the similarity and phylogenetic affinity of rock crawlers to the ice crawlers (suborder Grylloblattodea ).

COMMENTS: The new species, like all fossil Notoptera , lack the setal fan on the arolium, a feature synapomorphic for the modern, African species of Mantophasmatodea . like Raptophasma kerneggeri Zompro the new species lacks mesofemoral spination but otherwise has all of the features of the genus Adicophasma as characterized by Engel and Grimaldi (2004).

Zompro (2005) considered the holotype of A. spinosum as a nymph solely on its smaller body size. However, A. spinosum has fully developed genitalia indicative of an adult and has numerous significant features distinguishing it from R. kerneggeri as noted in the original description ( Engel and Grimaldi, 2004) and in the following key. Perhaps most notably the adult of Raptophasma is supposed to lack femoral spines, while they are exceptionally well developed in Adicophasma . Zompro’s statement to the contrary is entirely erroneous as the photograph in Engel and Grimaldi (2004) was not taken in the focal plane of the legs, yet the out-of-focus spines on at least one leg can nonetheless be discerned; similar, albeit slightly weaker, spines are obvious on A. grylloblattoides ( fig. 3 ). That the spines are indeed present in A. spinosum is evidenced by a new photograph of the holotype taken in the focal plane of the legs ( fig. 5 ), demonstrating that statements as to the absence of these structures are meant to mislead. Nymphal forms of rock crawlers can have spines in the latest of nymphal stages, but only when such features are similarly present in the adult (i.e., the latest nymphal stages acquire adult-like femoral spination). Thus, even if the holotype of A. spinosum were a nymph (which the terminal structures indicate is not the case), it could scarcely be considered conspecific with R. kerneggeri , which is supposed to lack such features entirely in the adult stage [as evidenced by the fact that the adult holotype male lacks such characters, as do, apparently, the paratypes; since spines in at least one paratype were entirely overlooked ( fig. 5 ), the series should be re-examined]. The same can be said for the dorsal profemoral carinae present in R. kerneggeri but lacking in Adicophasma , among other generic features (see Engel and Grimaldi, 2004). Most descriptions and figures of Baltic amber rock crawlers are inadequate (e.g., Zompro, 2001, 2005) as evidenced by the oversight of obvious features such as the presence/absence of femoral spination among paratypes and should not be relied upon. For more extensive descriptions and figures refer to Engel and Grimaldi (2004) and Grimaldi and Engel (2005). As the subfamily Raptophasmatinae and the family Ensiferophasmatidae are based entirely on plesiomorphies, they are herein newly synonymized with Mantophasmatidae (new synonymies). The Tyrannophasmatini is also suspicious and thus is only tentatively retained (table 1). Indeed, the recognition of multiple subfamilies or tribes within this remarkably small and relatively homogeneous family is unwarranted.

KEY TO BALTIC AMBER NOTOPTERA

1. Compound eyes occupying major portion of head; antennae shorter than body; arolium projecting beyond pretarsal claws...... 2

_ Compound eyes placed in posterior half of head; antennae several times longer than body; arolium not projecting beyond pretarsal claws (Genus Ensiferophasma).................... E. velociraptor Zompro

2. At least some femora with distinct, ventral spination; profemur without dorsal carinae (Genus Adicophasma )............... 3

_ Femora without spination; profemur with paired, dorsal, longitudinal carinae (Genus Raptophasma )..... R. kerneggeri Zompro

3. Mesofemur with distinct, ventral spination; pedicel elongate, at least three times longer than wide; cardo with two stout, apical spines; pronotum slightly longer than wide, meso- and metanota each wider than long; lateral margins of abdomen convex (i.e., abdomen roughly ovoid); compound eyes with large ommatidial facets (see Engel and Grimaldi, 2004)............................ A. spinosum Engel and Grimaldi

_ Mesofemur without ventral spination; pedicel short, about as long as wide; cardo without spines; all thoracic nota longer than wide and each of approximately equal size; abdomen elongate; compound eyes with smaller and more numerous ommatidial facets..................... A. grylloblattoides , n.sp.

Figures

Fig. 1. Photomicrograph of holotype of Adicophasma grylloblattoides Arillo and Engel, new species

Fig. 2. Illustration of holotype of Adicophasma grylloblattoides Arillo and Engel, new species

Fig. 3. Photomicrograph of holotype foreleg of Adicophasma grylloblattoides Arillo and Engel, new species (MCNA 10686).

Fig. 4. Illustration of head and pro- and mesothoracic segments of Adicophasma grylloblattoides Arillo and Engel, new species (MCNA 10686).

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 also present on the midlegs; see Engel and Grimaldi, 2004).