Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")

Abstract

The subfamily Platycraninae Brunner v. Wattenwyl, 1893 is polyphyletic in the traditional treatment. None of the genera attributed to the subfamily is closely related to the type-genus Platycrana Gray, 1835, which has already been suggested by morphological studies. The main key feature of Platycraninae traditionally used to characterize the subfamily, the remarkably large head and strongly enlarged genae ("cheeks"), is not true for Platycrana. Consequently, all other genera, the true "Palm Stick Insects" need to be removed from Platycraninae and are here accommodated in Megacraniinae subfam. nov.. Megacrania Kaup, 1871 is chosen as the type-genus of this new subfamily, because the name very well describes the main characteristic of this clade. Previous molecular studies have shown Megacraniinae subfam. nov. to be a member of the Old World family Phasmatidae s. str. (= Lanceocercata Bradler, 2001), which is characterized by the enlarged, laterally flattened and foliaceous or lanceolate cerci and by the male anal segment longitudinally split into two movable, interiorly dentate hemi-tergites that serve as a clasping apparatus to grasp the female abdomen during copulation and makes lacking a vomer. Neither character is true for Platycrana and Platycraninae sensu nov. respectively. Molecular studies have revealed Platycraninae sensu novo as the sister group of Phasmatidae s. str. (= Lanceocercata). The tribe Stephanacridini Günther, which has previously been without a subfamilar affiliation is transferred as a subordinate taxon of Platycraninae sensu novo. Platycraninae is characterized and distinguished from Phasmatidae s. str. (= Lanceocercata) by the small cylindrical cerci, simple anal segment and presence of a vomer in males, as well as the strongly keeled subgenital plate and remarkably elongated, filiform gonapophysis VIII of females, which greatly project over the apex of the abdomen. Characterizations are presented for Megacraniinae subfam. nov. as well as Platycraninae sensu nov. and its two tribes Platycranini and Stephanacridini. Lists of genera are provided for the three taxa. Three genera are removed from Platycraninae and Megacraniinae subfam. nov.: The Australian Echetlus Stål, 1875 is transferred to Phasmatinae: Acanthomimini and Redtenbacherus Özdikmen & Darilmaz, 2008 and Elicius Günther, 1935 are transferred to Lonchodidae: Necrosciinae. The male of Megacrania spina Hsiung, 2007 as well as the eggs of Megacrania brocki Hsiung, 2002, Erastus apalamnus Rehn, 1904, Ophicrania apterus (Redtenbacher, 1908), Ophicrania nigroplagiatus (Redtenbacher, 1908), Platycrana viridana (Olivier, 1792) and Macrophasma lyratus (Redtenbacher, 1908) are illustrated for the first time. Xenomaches moderata (Kirby, 1884) from the island of Ambon is synonymised with Platycrana viridana (Olivier, 1792) (syn. nov.).

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ZOOTAXA
ISSN 1175-5326 (print edition)
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Accepted by B. Mantovani: 8 Oct. 2020; published: 21 Dec. 2020 151
Zootaxa 4896 (2): 151–179
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4896.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:0BAD0251-42BC-4C88-BDDC-9622FD1F9F13
Megacraniinae—The Palm Stick Insects: A new subfamily of Old World
Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893
(Phasmatodea: “Anareolatae”)
FRANK H. HENNEMANN
Tannenwaldallee 53, 61348 Bad Homburg Germany.
�
hennemann@phasmatodea.com
Website: www.Phasmatodea.com
ZooBank: http://zoobank.org/651FFCA-271B-48A3-A58E-A30FDC739493
https://orcid.org/0000-0002-6547-4704
Abstract
The subfamily Platycraninae Brunner v. Wattenwyl, 1893 is polyphyletic in the traditional treatment. None of the genera
attributed to the subfamily is closely related to the type-genus Platycrana Gray, 1835, which has already been suggested
by morphological studies. The main key feature of Platycraninae traditionally used to characterize the subfamily, the
remarkably large head and strongly enlarged genae (“cheeks”), is not true for Platycrana. Consequently, all other genera,
the true “Palm Stick Insects” need to be removed from Platycraninae and are here accommodated in Megacraniinae
subfam. nov.. Megacrania Kaup, 1871 is chosen as the type-genus of this new subfamily, because the name very well
describes the main characteristic of this clade. Previous molecular studies have shown Megacraniinae subfam. nov. to
be a member of the Old World family Phasmatidae s. str. (= Lanceocercata Bradler, 2001), which is characterized by the
enlarged, laterally flattened and foliaceous or lanceolate cerci and by the male anal segment longitudinally split into two
movable, interiorly dentate hemi-tergites that serve as a clasping apparatus to grasp the female abdomen during copulation
and makes lacking a vomer. Neither character is true for Platycrana and Platycraninae sensu nov. respectively. Molecular
studies have revealed Platycraninae sensu novo as the sister group of Phasmatidae s. str. (= Lanceocercata). The tribe
Stephanacridini Günther, which has previously been without a subfamilar affiliation is transferred as a subordinate taxon
of Platycraninae sensu novo. Platycraninae is characterized and distinguished from Phasmatidae s. str. (= Lanceocercata)
by the small cylindrical cerci, simple anal segment and presence of a vomer in males, as well as the strongly keeled
subgenital plate and remarkably elongated, filiform gonapophysis VIII of females, which greatly project over the apex
of the abdomen. Characterizations are presented for Megacraniinae subfam. nov. as well as Platycraninae sensu nov.
and its two tribes Platycranini and Stephanacridini. Lists of genera are provided for the three taxa. Three genera are
removed from Platycraninae and Megacraniinae subfam. nov.: The Australian Echetlus Stål, 1875 is transferred to
Phasmatinae: Acanthomimini and Redtenbacherus Özdikmen & Darilmaz, 2008 and Elicius Günther, 1935 are transferred
to Lonchodidae: Necrosciinae. The male of Megacrania spina Hsiung, 2007 as well as the eggs of Megacrania brocki
Hsiung, 2002, Erastus apalamnus Rehn, 1904, Ophicrania apterus (Redtenbacher, 1908), Ophicrania nigroplagiatus
(Redtenbacher, 1908), Platycrana viridana (Olivier, 1792) and Macrophasma lyratus (Redtenbacher, 1908) are illustrated
for the first time. Xenomaches moderata (Kirby, 1884) from the island of Ambon is synonymised with Platycrana viridana
(Olivier, 1792) (syn. nov.).
Key words: Phasmatodea, Anareolatae, Phasmatidae, Lanceocercata, Megacraniinae; Platycraninae, Platycranini,
Stephanacridini, coconut stick insects, palm stick insects, new subfamily genera
Introduction
Considerable efforts have been made in the past fifteen years to re-arrange the taxonomy and reveal the natural re-
lationships within Old World Phasmatodea. Attempts have included morphological approaches (e.g. Bradler, 2001;
Hennemann & Conle, 2008; Hennemann et al., 2009, Hennemann et al., 2016), cladistic analyses of morphological
data (e.g. Tilgner, 2002; Bradler, 2009) as well as molecular-based phylogenetic studies (e.g. Whiting et al., 2003;
Buckley et al., 2010; Bradler et al., 2015; Robertson et al., 2018; Simon et al., 2019). Although the traditional clas-

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sification originally introduced by Günther (1953) gained broad acceptance by subsequent authors (e.g. Bradley &
Galil, 1977; Kevan, 1977; Kevan, 1982) the order was more recently considered to be lacking a phylogenetically
based classification (Bradler & Buckley, 2018). Many of the results obtained by recent studies did not corroborate
the traditional classification proposed by Günther (1953) and since then many inaccuracies have been uncovered
and clarified, these being important steps towards a natural classification. However, up to date still a good number
of relationships within the Old World Phasmatodea remain questionable.
One of the clades that still warrant clarification are the so-called “Coconut Stick Insects” or “Palm Stick Insects”
(traditionally the subfamily Platycraninae), a very charismatic subgroup of phasmatodeans that is known to feed ex-
clusively on various palms (Arecaceae) and screw palms (Pandanaceae) throughout the Oriental Region, southwest
Pacific Ocean and some islands of the Indian Ocean. Although, molecular-based phylogenetic studies (Buckley et
al., 2010) reveal Platycraninae as a member of the family Phasmatidae s. str. (= Lanceocercata Bradler, 2001) and
strongly support a monophyly of the clade, these studies have not included the eponymous type-genus Platycrana
Gray, 1835. Based on morphological characters a polyphyly of the traditional Platycraninae sensu Günther (1953)
has already been suggested by Hennemann & Conle (2008) and also by Bradler (2001; 2009), who presented il-
lustrations of the terminalia of males. This case well demonstrates the discrepancies that arise, when conclusions
on genera or higher ordinate taxa are being made without referring to the taxonomic type-species or type-genus
that these taxa were based on. In this particular case, the typical genus or type-genus of Platycraninae is not closely
related to all other genera that have traditionally been attributed to the subfamily. The main morphological key
features that characterizes this clade, the large head and remarkably enlarged genae (“cheeks”), is not true for the
type-genus of this clade. Moreover, not even the vernacular names “Coconut Stick Insects” or “Palm Stick Insects”
that are often used for Platycraninae, and which refer to the typical habits of these phasmatodeans, fit the type-genus
Platycrana. The anatomical adaptation of the head morphology, necessary to accommodate the massive mandibular
muscles needed for feeding on the hard palm leaves, is found in all genera traditionally attributed to Platycraninae
but not in Platycrana itself. Consequently and from a taxonomic point of view, a splitting of Platycraninae sensu
Günther is absolutely necessary and inevitable. To accommodate the genera removed from Platycraninae s. str., the
new subfamily Megacraniinae subfam. nov. is introduced. Megacrania Kaup, 1871 is chosen as the type-genus of
this new subfamily because the name very well describes the main characteristic of this clade, namely the remark-
ably enlarged head and genae.
The new subfamily is herein characterised and a list of genera included in Megacraniinae subfam. nov. is pro-
vided. In accordance to results of molecular studies by Buckley et al. (2010) and Bradler et al. (2015) the subfamily
is a member of Phasmatidae s. str. (= Lanceocercata), a well supported monophyletic clade of Old World Phasma-
todea, that is characterized by apomorphies such as the enlarged, laterally flattened and foliaceous or lanceolate
cerci and males having an anal segment that is longitudinally split into two movable, interiorly dentate hemi-tergites
that serve as a clasping apparatus to grasp the female abdomen during copulation, as well as lacking a vomer. The
subfamily Platycraninae sensu novo is here redefined, re-arranged and shown to comprise the tribe Stephanacri-
dini Günther, 1953, that has previously been revealed as the sister group of Phasmatidae s. str. (= Lanceocercata;
Buckley et al., 2010). Consequently, a sister group relationship can now be considered between Phasmatidae s. str.
(= Lanceocercata) and Platycraninae sensu nov.. This necessary taxonomic action is another much needed step
towards a meaningful re-classification and understanding of the Old World Phasmatodea.
Materials and methods
All specimens examined for this study are invariably dried and pinned. Insects and eggs were examined using
magnifying glasses with 4x and 10x magnification and a Russian MBC-10 binocular microscope. Measurements of
specimens and eggs were made to the nearest 0.1 mm using digital calipers. Photos of specimens were taken with a
Nikon D7000 camera equipped with a Nikon DX AF-S Macro 40 mm lens and a wireless Nikon SU-800 dual speed
light system. Background lightning was provided by a 18W 6000K LED panel light plate. The terminology and
abbreviations used for naming and marking of morphological parts of the genitalia, head and thorax of the insects
follows Beier (1968) and Bradler (2009). The terminology used to describe external and internal egg structures fol-
lows Clark-Sellick (1997).

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Abbreviations
All specimens here illustrated are within the authors private collection, abbreviated as “coll. FH”. Individual collec-
tion numbers and localities are provided in the captions of each figure. Below is a list of abbreviations that are used
for morphological parts and structures of the insects head, thorax and genitalia throughout all figures.
Ant = antenna
CE = compound eye
Cer = cerci
Cerv = cervix
CX = coxa
Ep = epiproct
Fi = fissure
Gap8 = gonapophysis 8
Gen = gena
Glo = glossa
Gpl = gonoplac
Gu = gula
LaCer = laterocervicalia
LabPa = labial palpus
LCF = lateral coronal fissure
MaxPa = maxillar palpus
Md = mandible
Me = mentum
MesoPl = mesopleura
MesoS = mesosternum
OpO = opercular organ
Pa = paraproct
PaGlo = paraglossa
PrO = praeopercular organ
ProBas = proabsisternum
ProN = pronotum
ProS = prosternum
S7, S8, ...= sternum 7, sternum 8, ...
Sg = subgenital plate
Sg (Po) = subgenital plate (poculum)
Sti8 = stigma 8
SubMe = submentum
T7, T8, ...= tergum 7, tergum 8, …
TP = thorn pads
Vo = vomer
Results
Megacraniinae subfam. nov.
Type-genus: Megacrania Kaup, 1871: 38.
Platycraniae Brunner v. Wattenwyl, 1893: 97 (in part – not the type-genus Platycrana Gray, 1835).
Redtenbacher, 1908: 368 (in part). [Phibalosomini (II. Sectio Platycraniae]
Platycraniinae Günther, 1953: 557 (in part). [Incorrect spelling based on misspelled Platycrania, Westwood, 1859]
Platycraninae Bradley & Galil, 1977: 190 (in part).

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Otte & Brock, 2005: 33 (in part).
Hennemann & Conle, 2008: 24, 26, 57 (in part).
Acrophyllinae Kirby, 1904: 379 (in part).
Cladoxerinae Karny, 1923: 237 (in part).
Diagnosis (♂, ♀): Small to medium-sized (body length 41.5–132.0 mm) often very colourful Phasmatidae, form
ranging from slender to fairly robust; body sub-cylindrical. ♂♂ apterous (Fig. 3D), brachypterous or with well
developed alae (Figs. 1B–C, 2A, 3A–C), ♀♀ apterous, brachypterous (Figs. 2B, D) or with moderately developed
alae (Figs. 1A, 2C). Sexual dimorphism moderate. Head conspicuously enlarged, notably broader and longer than
prothorax and flattened dorsoventrally, vertex smooth and with a distinct longitudinal postocular furrow laterally
(LCF); genae strongly enlarged (Fig 4A–J). Eyes more or less decidedly displaced towards the dorsal surface of
the head (Fig. 4A–J). No ocelli. Gula large and covering > ½ of cervical region (Fig. 5A–B). Lacinia usually with
three terminal teeth. Antennae moderately slim to robust and often perlamorph, distinctly segmented and at best
equal in length to profemora (much shorter than profemora in ♀♀); consisting of less than 30 segments. Two basal
antennomeres flattened dorsoventrally, the scapus moderately broadened. Pronotum with well developed, often very
prominent defensive glands at anterolateral corners; these lateral directed (Figs. 4A–J). Mesothorax > 2x longer than
prothorax; parallel-sided and more or less decidedly compressed dorsoventrally; mesonotum usually more or less
flattened, mostly smooth to sparsely granulose (Fig. 4G) but may be tuberculose (Fig. 4A) to spinulose in certain
taxa (Fig. 4C), with a fine medio-longitudinal carina. Tegmina flat and spatulate; in winged taxa notably shorter than
alae. Anal fan of alae transparent, plain grey, pink or orange; only in one known case with a darker outer margin.
Median segment longer than metanotum. Abdomen excluding median segment considerably longer than head and
complete thorax combined. Abdominal segments II–VI longer than wide. No praeopercular organ on abdominal
sternum VII of ♀♀ (Fig.. 8E). Terminalia of ♂♂ (Figs. 6, 7): Anal segment tectiform and split longitudinally to form
two movable hemi-tergites that are dorsally connected by a narrow fissure (FI, Figs. 7B, E, G–J); interior surface of
hemi-tergites with various specializations, either with paired medio-ventral directed thorn-pads (Fig. 6C), or with
often asymmetrical rows of teeth or dentate ridges that are directed medially against each other (Figs. 6A, 6B, 7B,
7E). No vomer (Figs. 6A, C). Paraprocts variable. Poculum (= subgenital plate) small, rather flat to moderately
convex and scoop-shaped (Figs. 7A, C, D, F). Terminalia of ♀♀ (Fig. 8): Subgenital plate at best slightly projecting
beyond but mostly not reaching to apex of abdomen, obtusely convex to moderately keeled longitudinally; often
with a knob-, wart-, or transversely ridge-like basal swelling (here referred to as “opercular organ”, Figs. 8A–E).
Gonapophysis VIII more or less equal in length to gonapophysis IX, gonoplac variable in size, sometimes notably
enlarged (Fig. 8A) and occasionally fused with gonapophysis IX; all hidden within subgenital plate and not reach-
ing to apex of abdomen. Gonangulum present but small. Cerci in both sexes compressed laterally, more or less
enlarged and elongated, foliaceous or lanceolate in shape (Figs. 6–8). All femora and tibiae trapezoidal in cross-
section; tibiae usually less decidedly carinate. Tibiae without an area apicalis. Profemora somewhat constricted and
strongly curved in basal portion (Figs. 4A–E); dorsal carinae very slightly nearing, the posterodorsal carina at best
very slightly raised sub-basally and occasionally minutely denticulate. Medioventral carina of all femora midways
on ventral surface and often set with a variable number of small spines or teeth. Extrimities otherwise unarmed.
Basitarsi slender, not lobed dorsally.
Eggs (Figs. 12A–H): Medium-sized to very large, more or less alveolar and /or polygonal; capsule distinctly
longer and higher than wide. Dorsal surface often strongly convex. Polar area mostly more or less distinctly in-
dented. Capsule surface prominently sculptured, being granulose, tuberculose, rugulose, coriaceous or punctured;
usually with at least weak irregular longitudinal ridges and/or keels. Micropylar plate small, less than ½ the length
of capsule and notably displaced towards the posterior end of capsule; shape more or less rhomboidal. Micropylar
cup placed roughly in centre of plate. Internal micropylar plate closed. Operculum strongly raised and conical or
peg-like.
Comments: The subfamily Platycraninae was originally established by Brunner v. Wattenwyl (1893: 97) as
Platycraniae to include the genera Platycrana Gray, 1835, Graeffea Brunner v. Wattenwyl, 1868 and Arrhidaeus
Stål, 1877 (= Ophicrania Kaup, 1871). Platycrana is the genus-eponymum and hence the type-genus. Subsequent
authors (e.g. Günther, 1953; Bradley & Galil, 1977) traditionally characterized the subfamily mainly by the mor-
phological character combination “Cheeks broader than eye” and “Wings, but the tegmina in particular, shortened
or lacking”.

MEGACRANIINAE—THE PALM STICK INSECTS: A NEW SUBFAMILY Zootaxa 4896 (2) © 2020 Magnolia Press · 155
FIGURE 1. Habitus of of Megacrania Kaup, 1871 species (Megacraniinae subfam. nov.): A. Megacrania phelaus (Westwood,
1859) ♀: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-7]; B. Megacrania phelaus (Westwood, 1859)
♂: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-8]; C. Megacrania spina Hsiung, 2007 ♂: West
Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1083-1].

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FIGURE 2. Habitus of Graeffea Brunner v. Wattenwyl, 1868 and Ophicrania Kaup, 1871 species (Megacraniinae subfam.
nov.): A. Graeffea meridionalis Günther, 1932 ♂: Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH,
No. 1195-1]; B. Graeffea crouanii (Le Guillou, 1841) ♀: Melanesia, Tonga, Tongatapu [coll. FH, No. 0204-1]; C. Ophicrania
nigroplagiatus var. (Redtenbacher, 1906) ♀: Philippines, Luzon Island [coll. FH, No. 0206-1]; D. Ophicrania apterus var.
(Redtenbacher, 1908) ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0222-1].
However, Platycrana Gray, 1835 (Figs. 9–10) the type-genus of Platycraninae, differs fundamentally from all
other genera currently attributed to the subfamily in numerous morphological characteristics and does not even ex-
hibit the aforementioned main diagnostic characteristic of the subfamily, the distinctly enlarged genae (or “cheeks”).
Hence, the subfamily in its present recognition is obviously polyphyletic. This has already been pointed out and
illustrated by Bradler (2001: 181, fig. 2) and Bradler (2009: 63, fig. 33), who studied the external genitalia of ♂♂,
as well as by Hennemann & Conle (2008: 24, 26), who in their distinguishing key to the subfamilies and tribes of
Phasmatidae s. str. (= Lanceocercata Bradler, 2001) referred to “Platycraninae” as comprising all genera attributed
to the subfamily at that time except for the type-genus Platycrana itself. In addition to stating that Platycrana was
not closely related to the remaining genera of Platycraninae in its traditional recognition, Hennemann & Conle

MEGACRANIINAE—THE PALM STICK INSECTS: A NEW SUBFAMILY Zootaxa 4896 (2) © 2020 Magnolia Press · 157
(2008: 57) suggested close relation between Platycrana and the tribe Stephanacridini Günther, 1953 (Fig. 11). Bra-
dler (2001: 181, fig. 2; 2009: 63, fig. 33) provided figures of the genitalia of ♂♂ of Platycrana and two members
of Megacraniinae subfam. nov., that clearly illustrate the fundamental differences in genital morphology. In ♂♂
of Platycrana the anal segment is flat and entire with the posterior margin just gently indented and the vomer is
well developed as a roughly triangular, strongly sclerotised plate that bears a terminal hook (Fig. 6D). The cerci
are small, cylindrical and do not show any specializations. All other genera formerly attributed to Platycraninae,
and here comprised in Megacraniinae subfam. nov., have the anal segment more or less tectiform and split longi-
tudinally into two movable hemi-tergites that bear variable dentate specializations on their interior surface to form
a clasping apparatus analogous to that of other Phasmatidae s. str. (= Lanceocercata, Figs. 6–7). Also the genital
morphology of ♀♀ of these two clades shows fundamental differences. In Platycrana the subgenital plate is much
elongated and projects considerably beyond the apex of the abdomen, there is a praeopercular organ on abdominal
sternum VII but no analogous structure at the base of the subgenital plate as in Megacraniinae subfam. nov., the
cerci are small and cylindrical, and the gonapophysis VIII are extremely elongated, filiform and project greatly
beyond the apex of the abdomen (Figs. 10A–C). In all other genera formerly attributed to Platycraninae, and here
comprised in Megacraniinae subfam. nov., the subgenital plate hardly projects over the apex of the abdomen and
bears an opercular organ (OpO) that is formed by a knob-, wart-, or transversely ridge-like swelling at the base
(Figs. 8A–E) and serves as an anchorage for the ♂♂ clasping apparatus during copulation, the cerci are laterally
compressed, more or less elongated and foliaceous or lanceolate (Figs. 6–8), and the gonapophysis VIII are not
notably elongated and fully hidden within the subgenital plate. Furthermore, the main key feature of Platycraninae
sensu Günther, 1953, the strongly enlarged genae (“cheeks”) only matches for the genera here comprised in Mega-
craniinae subfam. nov.. These have the genae strongly enlarged to accommodate the massive mandibular muscles
necessary for feeding on the hard leaves of various palms (Arecaceae) e.g. coconut (Cocos nucifera) or screw palms
(Pandanus spp., Pandanaceae). The head in these taxa is remarkably large, flattened dorsoventrally and the eyes are
more or less distinctly displaced towards the dorsal surface of the head capsule (Fig. 4). Moreover, the head capsule
bears a distinct longitudinal postocular furrow here referred to as the lateral cervical furrow (LCF, Fig. 4). In con-
trast, Platycrana has a fairly usual phasmatodean head, that is ovoid and somewhat widened towards the posterior
with the vertex roundly convex and the eyes placed on the lateral surfaces of the head capsule (Figs. 10H–J). Finally,
also the eggs of Platycrana are unlike those of all other genera of Platycraninae sensu Günther, 1953 (here com-
prised in Megacraniinae subfam. nov.), being ovoid with a flat operculum that bears a small knob-like capitulum in
the centre and having a large micropylar plate which exhibits two conspicuous lateral hook-like expansions (Figs.
12J–K). All genera in Megacraniinae subfam. nov. have more or less alveolar and polygonal eggs that bear a large,
conical to peg-like operculum and have a fairly small mostly rhomboidal micropylar plate (Figs. 12A–H).
As already suggested by the distinctive genital morphology of ♂♂ in particular, not only phylogenetic ap-
proaches based on morphological characters (e.g. Bradler, 2009) but also molecular-based phylogenetic studies
(Buckley et al., 2009; Bradler et al., 2015) have frequently revealed Megacraniinae subfam. nov. (in all men-
tioned studies referred to as “Platycraninae”) as a subordinate clade of Phasmatidae s. str. (= Lanceocercata). Thus
Platycrana, and the subfamily Platycraninae sensu novo consequently, are here removed from Phasmatidae s. str.
(= Lanceocercata), and based on morphological characters shown to be closely related to Stephanacridini (→ see
discussion of Platycraninae below). Molecular studies (Buckley et al., 2009) have recovered Stephanacridini as the
sister group of Phasmatidae s. str. (= Lanceocercata) and in removing Platycraninae from this clade, Stephanacridini
becomes a subordinate taxon of Platycraninae sensu nov.. Consequently and in accordance to the results of Buckley
et al. (2010), Platycraninae instead of Stephanacridini can now be considered as the sister group of Phasmatidae s.
str. (= Lanceocercata; Fig. 13). A detailed discussion of Platycraninae sensu novo and the arrangement of tribes and
genera within the subfamily is presented below.
Given that Platycrana, the type-genus of Platycraninae, is not closely related to any of the other genera tradi-
tionally attributed to that subfamily a new subfamiliar name is necessary to accommodate these genera. In order
to emphasize the main key feature of this subfamily, i.e. the remarkably enlarged head and genae, Megacraniinae
subfam. nov. is here introduced with Megacrania Kaup, 1871 as the type-genus. This new subfamiliar name, a
combination of the Greek μέγας or mega (= large) and κρανίον or cranium (= skull), is chosen because it very well
describes the main key characteristic that distinguishes this subfamily from all other related taxa. The vernacular
name “Palm stick insects” is here preferred to “Coconut Stick Insects”, as used by Bradler et al. (2015), because
members of this subfamily are not exclusively feeding on coconut palms but are known to feed on a variety of dif-

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ferent Arecaceae (e.g. Caryota spp., Cycas spp., Cocos nucifera, Trachycarpus spp., Metroxylon spp.) and screw
palms of the family Pandanaceae.
FIGURE 3. Habitus of Ophicrania Kaup, 1871 species ♂♂ (Megacraniinae subfam. nov.): A. Ophicrania vittipennis (Stål,
1875): Philippines, Leyte Island, Mahaplag [coll. FH, No. 0726-12]; B. Ophicrania capito (Westwood, 1859): Borneo, Sabah,
Crocker Range, Mt. Trus Madi 1200m [coll. FH, No. 0039-2]; C. Ophicrania sp. nov.: Borneo, Kalimantan, Malinau [coll. FH,
No. 0832-1]; D. Ophicrania sagittarius Bresseel & Bushell, 2009: Philippines, Mindanao Island, Agusan del Sur, Rosario [coll.
FH, No. 1131-1].

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FIGURE 4. Details of head and prothorax, A, B, D-G dorsal view, C dorsolateral view, H-J lateral view (Megacraniinae subfam.
nov.): A. Megacrania brocki Hsiung, 2002 ♀: Indonesia, Prov. Maluku, Buru Island [coll. FH, No. 1012-3]; B. Megacrania sp.
nov. ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-2]; C. Megacrania spina Hsiung, 2007 ♂: West Papua,
Prov. Papua Barat, Fak-Fak [coll. FH, No. 01083-1]; D. Erastus apalamnus Rehn, 1904 ♂: Indonesia, Prov. Maluku, Obi Island
[coll. FH, No. 1212-5]; E. vittipennis (Stål, 1875): Philippines, Leyte Island, Mt. Balocaue [coll. FH, No. 0726-8]; F. Ophicrania
capito (Westwood, 1859): Borneo, Sabah, Crocker Range [coll. FH, No. 0039-1]; G. Graeffea meridionalis Günther, 1932 ♂:
Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH, No. 1195-1]; H. Erastus apalamnus Rehn, 1904 ♀:
Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5]; J. Megacrania phelaus (Westwood, 1859) ♂: captive reared from
Solomon Islands, Malaita Island [coll. FH, No. 0174-10]. Arrow indicates the opening of the prothoracic defensive glands.

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FIGURE 5. Details of head and prothorax in ventral view (Megacraniinae subfam. nov.). A. Graeffea meridionalis Günther,
1932 ♂: Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH, No. 1195-1]; B. Erastus apalamnus Rehn,
1904 ♀: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5].
It appears noteworthy, that among Phasmatidae s. str. (= Lanceocercata) members of Megacraniinae subfam.
nov. show two striking morphological features that otherwise are exclusive to genera of the New Zealand clade
Acanthoxylini Günther, 1953. The opercular organ (OpO) is a conspicuous swelling at the base of the ♀♀ subgenital
plate that serves as an anchorage for the ♂♂ clasping apparatus during copulation. It varies considerably in size,
shape and sculpturing among the species and genera of Megacraniinae subfam. nov. ranging in shape from knob-
, over wart-like to forming an obtuse transverse ridge. While this structure is very prominent in e.g. Megacrania
Kaup, 1871 (Figs. 8C–E), Erastus Redtenbacher, 1908 (Fig. 8A) and Graeffea Brunner v. Wattenwyl, 1868 (Fig.
8B), it is just weakly developed to almost obsolete in Ophicrania Kaup, 1871. Being analogous to the structure in
Acanthoxylini it is seen to be similarly variable in this latter clade (see Buckley, Myers & Bradler, 2014: 460, fig.
4). The presence of an opercular organ could be a synapomorphy of Megacraniinae subfam. nov. + Acanthoxylini,
but further evaluation is needed to confirm this assumption. Another possible synapomorphy of these two clades is
exhibited by the eggs. In contrast to all other members of Phasmatidae s. str. (= Lanceocercata), which show a true
capitulum, the eggs of Megacraniinae subfam. nov. and Acanthoxylini have a more or less prominently raised and
conical or peg-like operculum (Figs. 12A–H), which may be as high as more than half the length of the egg capsule
in some extreme cases (e.g. some species of Ophicrania). Moreover, eggs of both clades are more or less polygonal
with the dorsal surface bulgy longitudinally and the polar-area more or less prominently indented. It shall be men-

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tioned that there have been contrary definitions of the opercular structures of genera of Megacraniinae subfam. nov.
in the past. According to Clark Sellick (1988: 278) the eggs of Megacrania and Ophicrania have “raised spongy
opercula” but according to Clark Sellick (1997: 118, figs. 71–72) capitula are present in the two genera. Since a true
capitulum is defined as a structure that is easily detached from the operculum without damage to the latter and with
its loss not having negative influence on the hatching rate (Clark, 1976; Clark Sellick, 1988), there is no capitulum
in Megacraniinae subfam. nov..
A detailed differentiation and comparison between Platycrana Gray, 1853 and Megacraniinae subfam. nov. is
presented in table 1 below.
FIGURE 6. Details of terminalia of ♂♂: A. Megacrania spina Hsiung, 2007, ventral view; B. Megacrania spina Hsiung, 2007,
caudal view of anal segment (tergum X); C. Ophicrania conlei Gottardo, 2011, ventral view; D. Platycrana viridana (Olivier,
1792), ventral view.

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FIGURE 7. Terminalia of ♂♂ of Megacraniinae subfam. nov.: A. Megacrania spina Hsiung, 2007, lateral view: West Papua,
Prov. Papua Barat, Fak-Fak [coll. FH, No. 1083-1]; B. Megacrania spina Hsiung, 2007, dorsal view: West Papua, Prov. Papua
Barat, Fak-Fak [coll. FH, No. 1083-1]; C. Megacrania spina Hsiung, 2007, ventral view: West Papua, Prov. Papua Barat, Fak-
Fak [coll. FH, No. 1083-1]; D. Megacrania sp. nov., lateral view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-
3]; E. Megacrania sp. nov., dorsal view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-3]; F. Graeffea leveri
(Günther, 1937), lateral view: captive reared from Solomon Islands, Russell Island NW of Guadalcanal [coll. FH, No. 0635-1];
G. Ophicrania capito (Westwood, 1859), dorsal view: Borneo, Sabah, Crocker Range, Mt. Trus Madi 1200m [coll. FH, No.
0039-2]; H. Ophicrania conlei Gottardo, 2011, dorsal view: Philippines, Mindoro Island, Mt. Halcon [coll. FH, No. 0611-1]; J.
Ophicrania sp. nov.: Borneo, Kalimantan, Malinau [coll. FH, No. 0832-1].
Genera excluded from Megacraniinae: The Australian Echetlus Stål, 1875 (Type-species: Bacillus peris-
thenes Westwood, 1859: 13, pl. 7: 1) was included in former Platycraninae by Günther (1953: 557) and has since
been retained in the subfamily by all subsequent authors. Brock & Hasenpusch (2007: 74) and Hennemann & Conle
(2008: 28) doubted Echetlus to be a member of Platycraninae sensu Günther and stated it was likely to belong in
Phasmatinae. Indeed, Echetlus is obviously not at all closely related to either Platycrana, and the subfamily Platy-
craninae sensu nov. respectively, nor to any of the genera of Megacraniinae subfam nov.. Consequently, since the
genus does not have any of the characteristics and can hence not be incorporated in either Platycraninae sensu nov.

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or Megacraniinae subfam. nov., it is here provisionally transferred to Phasmatinaea: Acanthomimini. Morphologi-
cal features such as the slender and stick-like habitus, the characteristic profemora which have the anterodorsal ca-
rina strongly raised and serrate and are triangular in cross-section, the elongate and cylindrical head as well as short
and stocky antennae that are considerably shorter than the profemora and also the geographic distribution in Western
Australia suggest close relation to genera currently in Acanthomimini. Hennemann & Conle (2008: 28) also pointed
out that the the Brazilian species attributed to Echetlus by Zompro (2004) is not congeneric and is a member of the
New World Diapheromeridae: Diapheromerinae.
FIGURE 8. Terminalia of ♀♀ of Megacraniinae subfam. nov.: A. Erastus apalamnus Rehn, 1904, lateral view (with egg):
Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5]; B. Graeffea crouanii (Le Guillou, 1841), lateral view: Melanesia,
Tonga, Tongatapu [coll. FH, No. 0204-2]; C. Megacrania sp. nov., lateral view: Papua New Guinea, Morobe Prov., Aseki [coll.
FH, No. 1014-2]; D. Megacrania phelaus (Westwood, 1859), lateral view: Solomon Islands, Maramasike Island prope Malaita
[coll. FH, No. 0174-7]; E. Megacrania tsudai Shiraki, 1933, ventral view: Taiwan, Pindang County, Gio-Pon [coll. FH, No.
0493-1]; F. Megacrania phelaus (Westwood, 1859), dorsal view: Solomon Islands, Maramasike Island prope Malaita [coll. FH,
No. 0174-7].
Redtenbacherus Özdikmen & Darilmaz, 2008 was introduced as a replacement name for the preoccupied Er-
nodes Redtenbacher, 1908 (Type-species: Ernodes sumatranus Redtenbacher, 1908: 374). In discussing Redten-
bacherus and transferring all species except for the type-species to Lopaphus Westwood, 1859 (subfamily Necro-
sciinae), Seow-Choen (2018: 450) commented “This species appears to be a Lopaphus species […]. The antennae
appear damaged, and therefore artificially shortened. It is likely to be proven to belong to that genus when more
information is available.”. Indeed, R. sumatranus strongly resembles certain species of Lopaphus and the species
has none of the characteristic features of either Platycraninae sensu nov. or Megacraniinae subfam. nov.. If taking

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broken antennae into account in the unique holotype, the slender and cylindrical body and stick-like habitus, entirely
unarmed extremities, triangular cross-section of the profemora, which have the anterodorsal carina strongly raised,
morphology of the head and genitalia clearly place this genus in the subfamily Necrosciinae. Consequently, and
since it cannot be retained in either Platycraninae nor Megacraniinae, Redtenbacherus is here transferred to Necro-
sciinae.
TABLE 1. Differentiation and comparison of Megacraniinae subfam. nov. and Platycrana Gray, 1835
Megacraniinae subfam. nov. Platycrana
Body cross-section Sub-cylindrical, ± flattened dorsoventrally Oval (♀♀) to cylindrical (♂♂)
Head Dorsoventrally flattened & sub-cylindrical
in cross-section; vertex flat (Fig. 4H–J)
Strongly globose with vertex roundly con-
vex (Figs. 10G–J)
Genae Strongly enlarged (Figs. 4A–J) Widened towards posterior of head (Fig.
10H)
Eyes ± strongly displaced towards dorsal surface
of head capsule (Figs. 4A–J)
On lateral surfaces of head capsule (Fig.
10H)
Gula Large and expanding > ½ of cervical region
(Fig. 5)
No; only with a pair of small gular sclerites
Mesonotum (♀♀) ± flattened and angulate laterally; may be
granulose to spinulose
Convex longitudinally with lateral margins
deflexed, rounded and overlapping pleurae
Mesonotum (♂♂) ± flattened and angulate laterally; may be
granulose to spinulose
Convex longitudinally; strongly tuberculose
to spinose (Fig. 10G)
Profemora Dorsal carinae ± equal; trapezoidal in cross-
section
Posterodorsal carinae decidedly raised; sub-
triangular in cross-section
Meso- and metafemora Four outer carinae unarmed Four outer carinae dentate
Cerci Compressed laterally, carinate ± enlarged
and foliaceous or lanceolate (Figs. 6–8)
Cylindrical; small (Figs. 6D, 10A–F)
Gonapophyses VIII (♀♀) Short and hidden within subgenital plate
(Figs. 8A–E)
Strongly elongated and filiform, project-
ing much beyond apex of abdomen (Figs.
10A–C)
Subgenital plate (♀♀) Short; at best projecting over apex of abdo-
men by length of anal segment;
often with a knob- or wart-like basal swell-
ing
Elongate and prominently keeled longitudi-
nally; projecting beyond apex of abdomen
by considerably more than length of anal
segment (Figs. 10A–C)
Anal segment (♂♂) Tectiform and consisting of two movable
hemi-tergites, that show dentate specializa-
tions at interior surface that serve as a clasp-
ing apparatus (Figs. 6A–C, 7)
Flattened and entire (Fig. 10D)
Vomer (♂♂) No (Figs. 6A, C) Well developed; a sclerotised, triangular,
hook-like organ (Fig. 6D)
Poculum (♂♂) Scoop-shaped Convex and cup-shaped
Egg capsule Laterally compressed, ± alveolar or polygo-
nal; polar area indented (Figs. 12A–H)
Ovoid (Figs. 12J–K)
Micropylar plate of eggs Small, rhomboidal; notably displaced to-
wards polar end of capsule (Fig. 12A–H)
Large, cross-shaped and with two conspicu-
ous up-curving lateral extensions; placed
in centre of dorsal capsule surface (Figs.
12J–K)
Operculum of eggs Conical or peg-shaped (Fig. 12A–H) Flat with a central knob-like capitulum
(Figs. 12J–K)

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The genus Elicius Günther, 1935 (Type-species: Elicius microbasileus Günther, 1935: 16) was described from
a unique male from Mount Latimojong, South Sulawesi. Already Günther (1935: 16) was in doubt about the sys-
tematic position and relationships of this tiny species and placed it in Platycraninae with reservation. The author
commented that the position of the genus within Platycaninae was just as incoherent as if it had been placed in the
subfamily Necrosciinae. However, detailed examination of the holotype in the Museum für Naturkunde Berlin,
Germany (MNHU) proves that Elicius is not a member of Megacraniinae subfam. nov. (= Platycraninae sensu
Günther). This is seen by morphological features such as the ovoid head that has the large eyes positioned laterally,
the genae not notably enlarged and lacks a lateral cervical furrow, the small gula, tectiform but entire anal segment,
presence of a well-developed vomer, cylindrical cerci, and long antennae that are almost as long as the front legs.
These features place Elicius in the subfamily Necrosciinae to which it is here transferred. The enlarged head as well
as the smooth and shiny body surface resemble certain Wallacean members of Paranecroscia Redtenbacher, 1908
and the genital morphology is similar to e.g. Moritasgus Günther, 19035, both members of the Necrosciinae.
Natural History: Many papers have dealt with the biology (e.g. Franzman, 1974; Bedford, 1978; Preston-Maf-
ham, 1990; Rapp, 1995; Cermak & Hasenpusch, 2000), ecology, host plants and pest status (e.g. Baker, 2015; Deesh
et al., 2020), population dynamics and dispersal (Kobayashi et al., 2014; Kobayahshi et al., 2016) and chemical
defensives (e.g. Smith et al., 1979; Chow & Lin, 1986; Ho & Chow, 1993; Jones & Bulbert, 2020) of members of
this particular subfamily. Natural host plants comprise various palms of the families Arecaceae and Pandanaceae.
For example, the so-called “Peppermint Stick Insect” Megacrania batesii Kirby, 1896 is reported to feed on Panda-
nus tectorius, P. monticola and P. soloms-laubachii in coastal parts of North Queensland, Australia (Cermak & Ha-
senpusch, 2000). The closely related Megacrania tsudai Shiraki, 1933 also feeds on Pandanus tectorius in similar
habitats in Taiwan (Chow & Lin, 1986). Several species, including Graeffea crouanii (Le Guillou, 1841), G. minor
Brunner v. Wattenwyl, 1868, G. lifouensis Sharp, 1898, G. leveri (Günther, 1937) and Acanthograeffea denticulata
(Redtenbacher, 1908) feed predominantly on coconut (Cocos nucifera, Arecaceae) in their natural habitats. While
Acanthograeffea denticulata has also been recorded on a species of Pandanus in the Marianas (Muniappan, 2002),
Graeffea crouanii has been observed to feed on sago (Metroxylon sagu, Arecaceae; Bedford, 1978) and the grass
Miscanthus floridulus (Poaceae) in Fiji (Lever, 1947). In captivity in Europe various palms of the Arecaceae family
are frequently accepted as substitutes by species of the genera Graeffea Brunner v. Wattenwyl, 1868 and Ophicrania
Kaup, 1871. These include Dypsis lutescens, Howea forestiana, Trachycarpus fortunei, Phoenix canariensis and
Livistona australis but also bamboo (Phyllostachyus aurea, Poaceae) was readily taken by G. crouanii in captivity
(Bruno Kneubühler, pers. comm.). Although Megacrania tsudai causes considerable damage to Pandanus screw
palms in Taiwan (e.g. Chow & Lin, 1986) and M. batesii to the same host plants in coastal parts of Northeast Aus-
tralia (Cermak & Hasenpusch, 2000), these damages are not of economic importance. The so-called “Coconut Stick
Insect” Graeffea crouanii however, is for long known to defoliate coconut palms (Cocos nucifera) in many islands
of the South Pacific and to be of great economic importance. Since coconut palms are one of the most important
crops and the coconut industry is the most important agricultural commodity for many smaller islands throughout
the Pacific, it plays important roles in the livelihoods of people in terms of food and nutrition security and economic
aspects. Hence, a decrease in infestation of coconut palms by G. crouanii is mandatory for the survival and boost
of the coconut industry (Deesh et al., 2013). The worst damage is mostly seen on old trees that are at least 25 m in
height and in times of severe damage, frequently whole plants are defoliated and die (Swaine, 1969). For example,
Paine (1968) reported a severe outbreak on the island of Taveuni, Fiji in 1958–1959 and extending into 1961 with
over 500 acres of coconut palms affected of which at least 50% of the older fronds were completely defoliated
and almost 400 palms killed. Many species of Megacraniinae subfam. nov. are found in coastal regions, which is
mostly due to the distribution and occurrence of their host plants. Kobayashi et al. (2014) suggested a dispersal of
eggs through seawater to be a possible distributional mechanism of Megacrania tsudai and tested this hypothesis
in laboratory conditions. The tests provided support for this assumption because eggs were shown to have specific
characteristics that allow their survival when they float in the sea and that the hatching rates were not negatively
affected. The same seawater resistance and dispersal by floating was reported for Graeffea crouanii in the South
West Pacific (Swaine, 1969). Members of the subfamily are also characteristic for having well developed bilateral
prothoracic defensive glands, which have their openings at the anterior corners of the pronotum. From these the in-
sects can spray good amounts of a white, milky and often strongly smelling and irritating secretion towards potential
predators. The secretion does not effect human skin (Franzmann, 1974) but irritates the eyes on contact (Cermak
& Hasenpusch, 2000). In species of Megacrania the smell resembles peppermint (e.g. Franzmann, 1974) and in

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Graeffea leveri the smell is described as lemon-like (Bruno Kneubühler, pers. comm). Gas chromatography-mass
spectrometry (GC-MS) analyses of the defensive secretion of Megacrania tsudai in Taiwan have shown it’s major
component to be actinidine but also to contain boschniakine and two stereoisomeres of 1-acetyl-3-methylclopten-
tane (Ho & Chow, 1993).
Distribution: Oriental Region from Japan and Taiwan in the north over Peninsular Malaysia, Borneo and the
Philippines towards Wallacea, the Papuan subregion and NE-Australia, and throughout the South West Pacific in
Melanesia, southern Micronesia, New Caledonia and as far South East as French Polynesia. One species is endemic
to the Seychelles and one possibly extinct genus, Xenomaches Kirby, 1896, was described from Rodrigues Island,
both in the Indian Ocean.
Genera included:
1. Acanthograeffea Günther, 1932: 760. Type-species: Graeffea denticulata Redtenbacher, 1908: 371, by original
designation.
Distribution: Northwestern Pacific, Mikronesia (Marianas & Carolines: Chuuk Islands).
2. Davidrentzia Brock & Hasenpusch, 2007: 51. Type-species: Davidrentzia valida Brock & Hasenpusch, 2007:
51, figs. 93–97, by original designation.
Distribution: Norfolk Islands (Lord Howe Island).
3. Erastus Redtenbacher, 1908: 373. Type-species: Erastus galbanus Redtenbacher, 1908: 373, pl. 17: 3, by sub-
sequent designation of Zompro, 2004: 310.
Distribution: New Guinea & Wallacea (Obi Island & Buru Island).
4. Graeffea Brunner v. Wattenwyl, 1868: 46. Type-species: Graeffea purpuripennis Brunner v. Wattenwyl, 1868:
46, figs. 1 & 2 (= Graeffea crouanii (Le Guillou, 1841)), by subsequent designation of Kirby, 1904: 386.
= Alopus Newport, 1844: 288. Type-species Alopus coccophagus Newport, 1844: 288, pl. 14: 4 (= Graeffea
crouanii (Le Guillou, 1841)), by original monotypy. [Synonymised by Kirby, 1904: 386]
Comments: In the present recognition this genus is obviously polyphyletic. Several of the species, currently attributed to
Graeffea are not congeneric to the type-species and belong either in Erastus Redtenbacher, 1908, Ophicrania Stål, 1875 or
elsewhere. This concerns to all those species that are not distributed throughout the Pacific Ocean, why these localities are
in square brackets below. A revision of Graeffea at the species level is needed.
Distribution: Southwestern Pacific (Samoa; Marquesas; Fiji); New Caledonia; Loyalty Islands; Solomon Islands; Sey-
chelles. [Wallacea (Ambon); New Guinea; Sulawesi].
5. Megacrania Kaup, 1871: 38. Type-species: Platycrania phelaus Westwood, 1859: 113, pl. 27: 5, by subsequent
designation of Kirby, 1904: 385.
Distribution: Taiwan; Japan; Borneo; Wallacea (Obi, Buru & Key Islands); New Guinea; Solomon Islands; Bismarck
Archipelago (Admirality Islands); NE-Australia; Vanuatu (New Hebrides); Fiji.
6. Ophicrania Kaup, 1871: 38. Type-species: Ophicrania striatocollis Kaup, 1871: 38, by original monotypy.
= Apterrhidaeus Karny, 1923: 238. Type-species: Arrhidaeus apterus Redtenbacher, 1908: 378, by original
designation. [Synonymised by Günther, 1953: 557]
= Arrhidaeus Stål, 1875: 40, 85. Type-species: Necroscia styxius Westwood, 1859: 136, pl. 9: 3, by original
monotypy [Synonymised by Günther, 1953: 557]
= Arrhidacus Brunner v. Wattenwyl, 1893: 97. [Misspelling of Arrhidaeus Stål]
Distribution: Philippines; Borneo; Peninsular Malaysia; Wallacea (Kaioa Island & Buru Island); New Guinea.
7. Xenomaches Kirby, 1896: 470. Type-species: Bacillus incommodus Butler, 1876: 8, pl. 54: 4, by monotypy.
= Xenomaces Bradley & Galil, 1977: 191. [Misspelling of Xenomaches]
= Xenomachus Redtenbacher, 1908: 379. [Misspelling of Xenomaches]
Comments: The second species formerly attributed to the genus, X. moderata (Kirby, 1884) from the Wallacean island
of Ambon, is here shown to be misplaced and is synonymised with Platycrana viridana (Olivier, 1792) syn. nov. (→ see
comments on Platycrana Gray, 1835 below).
Distribution: Rodrigues Island.
[?] Apterograeffea Cliquennois & Brock, 2002: 388. Type-species: Apterograeffea reunionensis Cliquennois &
Brock, 2002: 388, figs. 1–8, by original designation.
Comments: In studying the origin of the Mascarene stick insects on a molecular basis Bradler et al. (2015: 4) found that
Apterograeffea appears not to be a member of Megacraniinae subfam. nov. (referred to under Plarycraninae or “coco-

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nut stick insects”). Although this genus has at least one of the same adaptions to feeding on hard-leaved palms, e.g. the
noticeably enlarged genae that are necessary for accommodating the strong mandibular muscles, molecular data place
Apterograeffea apart from the true Megacraniinae and within an unnamed “Mascarene clade”, why this adaption is cur-
rently regarded as a convergent evolution. Since the genus was not formally removed from the traditional Platycraninae by
Bradler et al. (2015), it is here provisionally retained in Megacraniinae until more comprehensive work on the Mascarene
stick insects and formal naming of suggested clades are made.
Distribution: Mascarenes (Mauritius & La Réunion).
Platycraninae Brunner v. Wattenwyl, 1893 sensu nov.
Type-genus: Platycrana Gray, 1835: 36.
Platycraniae Brunner v. Wattenwyl, 1893: 97 (in part – only the type-genus Platycrana Gray, 1835)
Redtenbacher, 1908: 368 (in part). [Phibalosomini (II. Sectio Platycraniae]
Platycraniinae Günther, 1953: 557. [Incorrect spelling based on misspelled Platycrania Westwood, 1859]
Platycraninae Bradley & Galil, 1977: 190 (in part). [Corrected spelling]
Otte & Brock, 2005: 33 (in part).
Acrophyllini Redtenbacher, 1908: 436 (in part).
Cladoxerinae Karny, 1923: 237 (in part).
Lanceocercata Bradler, 2001: 179 (in part).
Pharnaciini Günther, 1953: 555 (in part).
Bradley & Galil, 1977: 193 (in part).
Phasminae Karny, 1923: 240 (in part).
Günther, 1953: 554 (in part).
Phasmatinae Bradley & Galil, 1977: 192 (in part). [Corrected spelling of Phasminae Karny, 1923: 240]
Phryganistriinae Kirby, 1904: 358 (in part).
Diagnosis (♂, ♀): Moderately sized to very large Phasmatodea (body length 76.0–300.0 mm) and comprising some
of the world’s longest insects. Very strong sexual dimorphism; ♂♂ stick-like, form of ♀♀ ranging from slender
to very stocky and sometimes remarkably massive. Both sexes apterous, brachypterous or pterous; apterous or
brachypterous ♀♀ often with corresponding ♂♂ brachypterous or with fully developed alae. Body surface rang-
ing from smooth and slightly glossy to moderately sculptured; mesonotum und mesopleurae often tuberculose or
spinose. Head ovoid to sub-spherical; no ocelli. No gula, only two small gular sclerites. Antennae much shorter
than body; in ♀♀ shorter than in ♂♂. Mesothorax slender in ♂♂, often more or less swollen medially in ♀♀ or
either lateral margins of mesonotum or mesopleurae deflexed and/or dentate. Median segment variable in length
but mostly shorter than metanotum. Praeopercular organ on sternum VII of ♀♀ indistinct (Figs. 6C, 11D, 11F).
Terminalia of ♂♂: Anal segment simple and entire (Figs. 6D, 11H, 11K), at best slightly tectiform an with posterior
margin indented medially; ventral surface of posterior margin with paired, ventral directed thorn-pads (TP, Figs. 6D,
10F, 11J, 11L). External vomer well developed and sclerotised; mostly triangular in shape and with a single terminal
hook (Figs. 6D, 10F, 11J, 11L). Terminalia of ♀♀: Gonapophysis VIII strongly elongated, filiform and projecting
considerably beyond apex of abdomen (Figs. 10A–C, 11D–G); much longer than gonapophysis IX or gonoplacs.
No gonangulum present. Subgenital plate prominently keeled longitudinally and usually projecting considerably
beyond apex of abdomen; shape variable (Figs. 10A–C, 11D–G). Extremities with most or all carinae more or less
strongly dentate, serrate and/or occasionally with single lobes. Profemora trapezoidal in cross-section, compressed
and curved basally; the medioventral carina indistinct and roughly midways on ventral surface of profemur; the
posterodorsal carina moderately raised. Meso- and metafemora trapezoidal in cross-section. Tibiae without an area
apicalis. Basitarsus longer than the following two segments combined.
Eggs (Figs. 12J–M): Small to large, shape variable; ranging from simply ovoid over almost spherical to con-
siderably longer than high with the cross-section distinctly oval and capsule surrounded by a dorso-ventral bulge.
Surface of capsule ranging from smooth and glossy to strongly sculptured. Internal micropylar plate closed, no
median line. Operculum flat, capitulum present but not stalked.
Comments: The subfamily Platycraninae was based on the genus Platycrana Gray, 1835. As discussed in detail
above, the type-genus Platycrana is not closely related with all other genera that were traditionally attributed to
the subfamily. These genera are here removed from Platycraninae and accommodated in Megacraniinae subfam.

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nov.. As a result, Platycrana is the only genus that remains in Platycraninae sensu nov.. Close relation between
Platycraninae sensu novo and the tribe Stephanacridini Günther, 1953 has already been suggested by Hennemann &
Conle (2008: 57) and based on a prolific set of morphological characters this relation is supported herein. Common
characters include the lack of a gula (only two small gular sclerites present), small and more or less cylindrical cerci
of both sexes, a simple and entire anal segment (Figs. 10D, 11K) and well developed external vomer of ♂♂ (Figs.
6D, 10E–F, 11J, 11L), as well as the more or less prominently elongated subgenital plate and often enormously
elongated and filiform gonapophysis VIII of ♀♀, that considerably project beyond the apex of the abdomen (Figs.
10A–C, 11D–G). The eggs share a flattened operculum and knob-like central capitulum (Figs. 12J–M). All these
features are not only shared by Platycrana and Stephanacridini, but they also distinguish both clades from Megacra-
niinae subfam. nov. and the entire family Phasmatidae s. str. (= Lanceocercata). Consequently, Stephanacridini was
removed from Phasmatidae s. str. (= Lanceocercata) by Hennemann & Conle (2008) which has received strong sup-
port by subsequent phylogenetic approaches based on morphological data (Buckley et al., 2009) and molecular data
(Buckley et al., 2009; Robertson et al., 2018). These studies have moreover recovered Stephanacridini as the sister
group of Phasmatidae s. str. (= Lanceocercata) but Stephanacridini has since remained without a subfamiliar affili-
ation. The here revealed close relation to Platycrana places Stephanacridini as a subordinate taxon of Platycraninae
sensu nov., which consequently means that automatically Platycraninae, instead of Stephanacridini, should now be
considered as the possible sister taxon of Phasmatidae s. str. (= Lanceocercata). Platycraninae senso nov. comprises
two tribes, i.e. the monotypical Platycranini, which only contains its type-genus Platycrana, and Stephanacridini.
Simon et al. (2019) have obtained Xenophasmina Uvarov, 1940, an Indo-Chinese genus that currently is con-
sidered to be a member of the polyphyletic subfamily Xeroderinae, as the sister taxon of Phasmatidae s. str. (=
Lanceocercata). Moreover, these authors considered that Xenophasmina might be a member of Stephanacridini.
From a morphological point of view however, there is absolutely no support for this assumption. Although ♂♂
of Xenophasmina possess an entire anal segment and a well developed vomer, both sexes have notably enlarged,
dorsoventrally flattened and more or less foliaceous cerci, that remarkably resemble the cerci found throughout
Phasmatidae s. str. (= Lanceocercata) and ♀♀ lack the elongated and filiform gonapophysis VIII seen in Stephana-
cridini. Moreover, the eggs of Xenophasmina lack the knob-like central capitulum seen in Stephanacridini but have
a strongly raised opercular structure with a central hollow instead and the internal micropylar plate is open with a
distinct median line (closed in Stephanacridini). Other remarkable morphological characters, such as the notably
flattened head, laterally lobed abdominal terga VII–X of both sexes and notably deflexed, undulate and strongly
setose lower outer carinae of all legs in ♀♀ distinguish Xenophasmina from Stephanacridini. Hence, an inclusion of
Xenophasmina in future analyses appears desirable in order to uncover the relationships of the genus.
Distributuion: Taiwan, Peninsular Malaysia, Borneo, Philippines, Wallacea, New Guinea, Solomon Islands,
NE-Australia, Micronesia and Melanesia (Fiji, Tonga, Samoa, Vanuatu, New Caledonia & Western French Polyne-
sia).
Tribes included:
1. Platycranini Brunner v. Wattenwyl, 1893: 97.
2. Stephanacridini Günther, 1953: 555.
Keys to the tribes of Platycraninae
♀♀
1. Winged, alae well developed and reaching to abdominal segment V; lateral margins of mesonotum deflexed, rounded and over-
lapping mesopleurae; pronotum transverse; mesonotum < 3.5x the length of pronotum . . . . . . . . . . . . . . . . . . . . . Platycranini
- Apterous or brachpterous, alae at best covering median segment; lateral margins of mesonotum not deflexed; pronotum longer
than wide; mesonotum > 3.5x longer than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Stephanacridini
♂♂
1. Stocky insects; mesonotum < 3.5x longer than pronotum; pronotum quadrate ............................. Platycranini
- More slender and elongate insects; mesonotum > 4x longer than pronotum; pronotum longer than wide . . . . . .Stephanacridini
Eggs
1. Micropylar plate very large, almost as long as capsule; shape complex with two large hook-shaped lateral expansions (Figs.
12J–K) .................................................................................... Platycranini
- Micropylar plate small and < 2/3 the length of capsule; shape ± rhomboidal (Figs. 12L–M) . . . . . . . . . . . . . . . .Stephanacridini

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Tribe Platycranini Brunner v. Wattenwyl, 1893
(Figs. 6D, 9–10, 12J–K)
Type-genus: Platycrana Gray, 1835: 36.
Platycraniae Brunner v. Wattenwyl, 1893: 97
Otte & Brock, 2005: 13. [First use as Platycranini – noted that technically not needed]
Seow-Choen, 2018: 449.
Diagnosis (♂, ♀): Moderately sized (body length ♂♂ 80.0–90.0 mm, ♀♀ incl. subgenital plate 125.0–155.0 mm),
very stocky and massive (♀♀ in particular) bright green Platycraninae (Fig. 9). Both sexes with fully developed alae
that cover more than half of abdomen; anal fan plain transparent pink. Head strongly globose with vertex rounded
and smooth. Pronotum quadrate (♂♂) or transverse (♀♀). Mesothorax short and less than 3.5x longer than protho-
rax. Mesonotum smooth in ♀♀ with the lateral margins strongly deflexed, rounded and overlapping mesopleurae; in
♂♂ slender and prominently tuberculose or spinose (Fig. 10G). Mesopleurae simple (minutely tuberculose in ♂♂).
Median segment longer than metanotum. Abdominal tergites II–VII of ♀♀ with lateral margins noticeably deflexed;
II–VI wider than long. Subgenital plate of ♀♀ strongly keeled longitudinally, boat-shaped and projecting over apex
of abdomen by considerably more than length of anal segment (Fig. 10A). Vomer of ♂♂ with a ridge-like to conical
basal swelling (Figs. 6D, 10D, 10F).
Eggs (Figs. 12J–K): Large and ovoid; colour plain pale to mid brown. Entire capsule surface deeply and
densely punctured; otherwise with some obtuse bulges. Micropylar plate very large, almost as long as capsule
and of a complex shape with two large, hook-like lateral expansions and the polar portion very narrow and almost
reaching to posterior pole; entire plate slightly lowered. Outer margin distinctly marked by dark brown, the inner
portion sculptured like capsule. Micropylar cup small and just above narrow posterior portion of plate. Internal
micropylar plate with a very long and slender posterior extension. Chorion very thick (ca. 0.7–0.8 mm), comb-like
and consisting of tube-like, radially directed hollow compartments. Operculum dark reddish brown, almost circular
and with a slightly swollen, somewhat granulose circular bulge some distance off the outer margin; the central por-
tion impressed. Capitulum small, mushroom-like in shape; orange. Measurements [mm]: overall length 6.3–6.5,
capsule length 5.9–6.1, width 4.0–4.2, height 4.3–4.7, length of micropylar plate 4.9–5.1, width of micropylar plate
3.6–3.7.
Comments: This tribe only contains the monotypical type-genus Platycrana Gray, 1835, that is widely distrib-
uted throughout Wallacea and some of the more southern Philippine Islands. Although P. viridana (Olivier, 1792)
appears to be fairly abundant in certain localities, almost nothing is currently known about the habits and natural
food plants. Morphologically the massive, bright green ♀♀ resemble winged version of certain species of Macro-
phasma Hennemann & Conle, 2006 (tribe Stephanacridini), e.g. M. oreitrephes (Günther, 1929). Since the egg has
not yet been formally described, a detailed description and measurements are here presented. The description is
based on two examples from the Philippine Island of Panay in the authors collection (coll. FH, No. 0173-E).
A questionable Wallacean species attributed to the Mascarene Megacraniinae genus Xenomaches Kirby, 1896
(→ see above) by Redtenbacher (1908: 379) and since retained in that genus by subsequent authors is not a member
of Xenomaches nor the subfamily Megacraniinae subfam. nov.. Examination of the holotype ♂ of Xenomaches
moderata (Kirby, 1884) from the island of Ambon in Natural History Museum, London, U.K. (NHMUK) has recov-
ered this to be a strongly discoloured specimen of Platycrana viridana (Olivier, 1792). The pale yellow colour was
obviously caused by long-term storage in ethanol. This species is here synonymised with P. viridana (syn. nov.).
Distribution: Wallacea and Philippines.
Genus included:
1. Platycrana Gray, 1835: 36. (Type-species: Mantis viridana Olivier, 1792: 636, by subsequent designation of
Kirby, 1904: 385.
= Platycrania Burmeister, 1838: 581. [Subsequent misspelling of Platycrana]

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FIGURE 9. Habitus of Platycrana viridana (Olivier, 1792), Platycraninae: Platycranini Brunner v. Wattenwyl, 1893: A. ♀ dor-
solateral view: Kei Islands [coll. FH, No. 0173-3]; B. ♀ dorsolateral view: Kei Islands [coll. FH, No. 0173-6]; C. ♀ dorsolateral
view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-11]; D. ♀ dorsolateral view: Philippines, Panay Island, Mt.
Nangtud [coll. FH, No. 0173-11].

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FIGURE 10. Details of Platycrana viridana (Olivier, 1792), Platycraninae: Platycranini: A. Terminalia of ♀, lateral view: Kei
Islands [coll. FH, No. 0173-5]; B. Terminalia of ♀, dorsal view: Kei Islands [coll. FH, No. 0173-5]; C. Terminalia of ♀, ventral
view: Kei Islands [coll. FH, No. 0173-6]; D. Terminalia of ♂, lateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH,
No. 0173-13]; E. Terminalia of ♂, dorsal view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; F. Terminalia of
♂, ventral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; G. Head, pro- and mesothorax of ♂ in lateral
view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; H. Head, pro- and mesothorax of ♀ in lateral view: Kei
Islands [coll. FH, No. 0173-3]; J. Head, pro- and mesothorax of ♀ in lateral view: Kei Islands [coll. FH, No. 0173-6].

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FIGURE 11. Habitus and details of members of Platycraninae: Stephanacridini Günther, 1953: A. Macrophasma oreitrephes
(Günther, 1929) ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0107-2]; B. Hermarchus insignis (Kaup & Heyden,
1871) ♀: captive reared from ‘Australian Islands’ [coll. FH, No. 0144-13]; C. Eucarcharus sp. ♂: captive reared from Philip-
pines, Mindanao Island, Prov. Bukidnon, Mt. Apo [coll. FH, No. 0630-7]; D. Hermarchus insignis (Kaup & Heyden, 1871)
terminalia of ♀, lateral view: captive reared from ‘Australian Islands’ [coll. FH, No. 0144-13]; E. Hermarchus insignis (Kaup
& Heyden, 1871) terminalia of ♀, dorsal view: captive reared from ‘Australian Islands’ [coll. FH, No. 0144-13]; F. Hermarchus
insignis (Kaup & Heyden, 1871) terminalia of ♀, ventral view: captive reared from ‘Australian Islands’ [coll. FH, No. 0144-13];
G. Stephanacris draconius Hennemann & Conle, 2006 terminalia of ♀, dorsal view: Papua New Guinea, Morobe Prov., Wau
[coll. FH, No. 0377-2]; H. Hermarchus pythonius var. (Westwood, 1859) terminalia of ♂, lateral view: captive reared from Fiji
Islands [coll. FH, No. 0144-2]; J. Hermarchus pythonius var. (Westwood, 1859) terminalia of ♂, ventral view: captive reared
from Fiji Islands [coll. FH, No. 0144-2]; K. Stephanacris globiceps Redtenbacher, 1908 terminalia of ♂, dorsal view: West Pa-
pua, Arfak Mountains, Manokwari, Maibri village, 670m [coll. FH, No. 1000-1]; L. Phasmotaenia lanyuhensis Huang & Brock,
2001 terminalia of ♂, ventral view: Taiwan, Lanyuh Island [coll. FH, No. 0616-2].

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FIGURE 12. Eggs of Megacraniinae subfam. nov. (A-H), Platycraninae: Platycranini (J-K) and Platycraninae: Stephanacridini
(L-M): A. Megacrania batesii Kirby, 1896, dorsolateral view: NE-Australia, Queensland, Cape Tribulation [coll. FH, No.
0175-E]; B. Megacrania tsudai Shiraki, 1933, dorsolateral view: Taiwan, Pindang County, Gio-Pon [coll. FH, No. 0493-E]; C.
Megacrania brocki Hsiung, 2002 dorsolateral view: Indonesia, Prov. Maluku, Buru Island [coll. FH, No. 1012-E]; D. Erastus
apalamnus Rehn, 1904, dorsal view: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-E]; E. Erastus apalamnus Rehn,
1904, dorsal view: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-E]; F. Ophicrania nigroplagiatus var. (Redtenbach-
er, 1906), dorsolateral view: Philippines, Luzon Island [coll. FH, No. 0206-E]; G. Graeffea crouanii (Le Guillou, 1841), dor-
solateral view: captive reared from Melanesia, Fiji, Suva [coll. FH, No. 0204-E2]; H. Ophicrania apterus var. (Redtenbacher,
1908), dorsolateral view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0222-E1], J. Platycrana viridana (Olivier,
1792), dorsal view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-E1]; K. Platycrana viridana (Olivier, 1792),
lateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-E1]; L. Hermarchus pythonius var. (Westwood, 1859),
dorsolateral view: captive reared from Fiji Islands [coll. FH, No. 0144-E1]; M. Macrophasma lyratus (Redtenbacher, 1908),
dorsolateral: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1081-E].

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Tribe Stephanacridini Günther, 1953
(Figs. 11, 12L–M)
Type genus: Stephanacris Redtenbacher, 1908: 441.
Stephanacridini Günther, 1953: 555.
Bradley & Galil, 1977: 194.
Hennemann & Conle, 2004: 48.
Otte & Brock, 2005: 33.
Hennemann & Conle, 2008: 54, figs. 37, 43, 60–61, 80–82.
Pharnaciini Günther, 1953: 555 (in part).
Bradley & Galil, 1977: 193 (in part).
Otte & Brock, 2005: 32 (in part).
Diagnosis (♂, ♀): Variably sized often very large Platycraninae (body length ♂♂ 76.0–186.5 mm, ♀♀ incl. subgen-
ital plate 135.0–300.0 mm), shape very variable and ranging from very slender and stick-like to stocky and robust.
♂♂ apterous, brachypterous or with fully developed alae, ♀♀ mostly apterous but sometimes brachypterous with
alae at best covering median segment and hardly longer than tegmina. If alae present, anal fan in ♂♂ transparent, in
♀♀ colourful. Pronotum longer than wide. Mesothorax elongate and > 3.5x longer than prothorax; always slender
in ♂♂. Mesonotum and mesopleurae often armed; in ♀♀ lateral margins of mesonotum at best slightly widened and
mesopleurae occasionally strongly deflexed. Abdominal segments II–VI of ♀♀ longer than wide (quadrate only in
one species). All extrimities armed, occasionally with single enlarged lobes. Basitarsi slender or lobed.
Eggs (Figs. 12L–M): Variable in shape, size and capsule sculpturing. Micropylar plate small, less than 2/3 the
length of capsule and more or less rhomboidal in shape. Micropylar cup placed roughly in centre of plate.
Comments: Stephanacridini was originally established by Günther (1953: 555) as a tribe of the subfamily Phas-
minae (= Phasmatinae Bradley & Galil, 1977) and originally only contained the type-genus Stephanacris Redten-
bacher, 1908. The fundamental differences concerning to the morphology of the genitalia and eggs, that prove the
tribe cannot be a member of Phasmatidae s. str. (= Lanceocercata) were summarized and illustrated by Hennemann
& Conle (2008: 55), who transferred several misplaced genera from Pharnaciini Günther, 1953 and provided a list
of genera included in Stephanacridini. However, Stephanacridini has since remained without a definite affiliation to
a subfamily and is here attributed to Platycraninae sensu nov..
The two New Guinean genera Stephanacris and Macrophasma Hennemann & Conle, 2006 have been revised
by Hennemann & Conle (2006) and a revision of Phasmotaenia Návas, 1907 was presented by Hennemann & Conle
(2009). A review of Nesiophasma Günther, 1934 is in progress by the author and the remaining four genera are in-
tended to be reviewed in the future as well.
Distribution: Taiwan, Borneo, Philippines, Wallacea, New Guinea, Solomon Islands, NE-Australia, Microne-
sia and Melanesia (Fiji, Tonga, Samoa, Vanuatu, New Caledonia & Western French Polynesia).
Genera included:
1. Diagoras Stål, 1877: 66. Type species: Diagoras ephialtes Stål, 1877: 66, by original monotypy.
= Eustygera Brunner v. Wattenwyl, 1907: 186. Type species Eustygera godeffroyi Brunner v. Wattenwyl, 1907:
186, pl. 7: 2, by original monotypy. [Synonymised by Günther, 1932b: 756]
Distribution: NW-Pacific (Caroline Islands, Palau).
2. Eucarcharus Brunner v. Wattenwyl, 1907: 185. Type species: Lonchodes feruloides Westwood, 1859: 45, pl. 6:
5a–b, by subsequent designation of Günther, 1935b: 138.
Distribution: Philippines, Java.
3. Hermarchus Stål, 1875: 45. Type species: Phibalosoma pythonius Westwood, 1859: 73, pl. 12: 1 & 35: 3, by
subsequent designation of Kirby, 1904: 361.
Distribution: Melanesia (Fiji, Tonga, Vanuatu, New Hebrides, New Caledonia, Western French Polynesia, NE- Australia).
4. Macrophas
ma Hennemann & Conle, 2006: 3. Type species: Hermarchus biroi Redtenbacher, 1908: 445, by
original designation.
Distribution: New Guinea.
5. Nesiophasma Günther, 1934: 5. Type species: Nesiophasma eremothocus Günther, 1934: 6, fig. 1, by original
designation.

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= Mylothrus Günther, 1935a: 18. Type species: Mylothrus oligarches Günther, 1935a: 18, pl. 2: 12, by original
designation. [Synonymised by Günther, 1935b: 29]
Distribution: Sulawesi, Selayar Island S of Sulawesi, Sangir Island, Lesser Sunda Islands (Romang Island), New Guin-
ea.
6. Phasmotaenia Návas, 1907: 10. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication.
[replacement name for the preoccupied Taenionema Bolivar, 1906]
= Taeniosoma Bolivar, 1897: 31. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by monotypy.
[Praeoccupied by Taeniosoma Stimpson, 1857 (Nematoda)]
= Taenionema Bolivar, 1906: 393. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication.
[Replacement name for preoccupied Taeniosoma Bolivar, 1897 - preoccupied by Taenionema Banks, 1905
(Plecoptera)]
= Taeniophasma Uvarov, 1940: 379. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication.
[Homonym of Phasmotaenia Návas, 1907 & unnecessary replacement name for Taenionema Bolivar, 1906]
= Phasmotaenionema, Günther, 1933: 155. [Misspelling of Phasmotaenia Návas, 1907]
= Phasmatotaenionema, Bradley & Galil, 1977: 193. [Misspelling of Phasmotaenia Návas, 1907]
Distribution: Lanyuh Island SE of Taiwan, Philippines, Micronesia, Solomon Islands, New Guinea, Fiji.
7. Sadyattes Stål, 1875: 44, 88. Type species: Sadyattes borrii Stål, 1875: 88, by original monotypy.
Distribution: Borneo, Engano Island (near Sumatra).
8. Stephanacris Redtenbacher, 1908: 441. Type species: Stephanacris brevipes Redtenbacher, 1908: 441, pl. 21:
4a–b, by subsequent designation of Bradley & Galil, 1977: 194.
Distribution: New Guinea.
Discussion
The subfamily Platycraninae in its traditional composition (sensu Günther, 1953) is a good example for polyphyletic
clades within Phasmatodea, an insect order that stands out as one of the few orders that still lack a phylogenetically
based classification (Bradler & Buckley, 2018). Phylogenetic analyses based on morphological (e.g. Tilgner, 2002;
Bradler, 2009) and molecular data (e.g. Whiting et al., 2003; Buckley et al., 2010; Bradler et al., 2015; Robertson et
al., 2018; Simon et al., 2019) clearly show that the Phasmatodea is still lacking a meaningful higher-level classifi-
cation. Based on morphological characters the polyphyly of Platycraninae sensu Günther has already been pointed
out by Hennemann & Conle (2008) and was illustrated by Bradler (2001; 2009). All subsequent phylogenetic stud-
ies of molecular data have revealed the traditional Platycraninae as a monophyletic clade within the Old World
Phasmatidae s. str. (= Lanceocercata), but unfortunately none of these studies has included the nominate genus (or
type-genus according to taxonomic rules, see ICZN) Platycrana. This case very well demonstrates the discrepan-
cies that arise, when conclusions on higher ordinate taxa and clades are being made without referring to the taxo-
nomic type-species or type-genus that these taxa and clades are based on. In this particular case, the type-genus of
Platycraninae is not closely related to all other genera that have traditionally been attributed to that subfamily. The
main morphological key feature that traditionally characterized this clade, the large head and remarkably enlarged
genae (“cheeks”), is true for all genera included except for the type-genus Platycrana itself. Nor do the vernacular
names “Coconut Stick Insects” or “Palm Stick Insects” that are often used to refer to traditional Platycraninae fit
the type-genus Platycrana. In other words, the name Platycraninae cannot be used for the clade that is traditionally
referred to under that name because it does not include the type-genus. Therefore a new subfamily name is neces-
sary for the so-called “Palm Stick Insects” and Megacraniinae subfam. nov. is chosen, because it well describes
the main characteristics of this clade, namely the strongly enlarged head and genae. Although the practical use of
traditional taxonomic ranks is more and more, and with good cause, called into question by various recent workers,
the significance and inclusion of taxonomic type-species and type-genera is of utmost importance for correct and
meaningful naming of clades, no matter whether these are of traditional taxonomic rank or rank-free. Hence, it is
hoped that future phylogenetic studies will attach importance to including type-species and type-genera, which in
this case is the genus Platycrana.
Examinations within the framework of this study have shown that most of the genera comprised in Megacra-
niinae subfam. nov. require proper redefinitions and delimitations and are in need of revision at the species level.

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At least Erastus Redtenbacher, 1908, Graeffea Brunner v. Wattenwyl, 1868 and Ophicrania Stål, 1875 appear to be
polyphyletic in their current compositions and if all species currently included in each genus are taken into account.
For instance, species that are not distributed throughout the Pacific Ocean and attributed to Graeffea are obviously
not congeneric with the type-species and either belong in Erastus or Ophicrania. The author is aware of several as
yet undescribed species, two of which are here illustrated (Figs 3C, 4B, 7D, 8C) and many species, particularly in
Ophicrania are so far only known from a single sex. Moreover, Ophicrania may contain some as yet unrevealed
synonymies but the strong sexual dimorphism and often striking differences in colouration between ♂♂ and ♀♀ of
individual species make correct matching of sexes fairly difficult without having at hand specimens from the same
locality and collecting event. Thus, much further work is needed for understanding the taxonomy and systematics of
Megacraniinae subfam. nov., which however should not exclusively aim on the descriptions of new taxa but should
also include re-describing the known taxa. It is hoped that more fresh material from throughout the distributional
range of the “Palm Stick Insects” will become available in the future for adding to our knowledge of this very dis-
tinctive and interesting clade of Old World Phasmatodea.
FIGURE 13. Supposed relationships between Phasmatidae s. str. (= Lanceocercata), including the subordinate Megacraniinae
subfam. nov., and the sister group Platycraninae sensu nov. with the two contained tribes Platycranini and Stephanacridini. The
tree shown is merely meant to visualize the estimated relationships here suggested. It does not restrict to defining apomorphies
for each clade but also uses key-features that are helpful for distinction. Although the results of Robertson et al. (2018, Fig. 6)
and Simon et al. (2019, Fig. 2), both based on molecular data, are incorporated the above tree cannot be interpreted as a strict
phylogenetic cladogram.

MEGACRANIINAE—THE PALM STICK INSECTS: A NEW SUBFAMILY Zootaxa 4896 (2) © 2020 Magnolia Press · 177
Acknowledgements
The author wants to thank the following curators for access to the collections of the corresponding museums and
institutions and/or providing information and pictures: Judith Marshall & Dr. George Beccaloni (Natural History
Museum London, U.K), Dr. Susanne Randolf, Dr. Ulrike Aspöck and Dr. Harald Bruckner (Naturhistorisches Mu-
seum, Vienna, Austria), Dr. Michael Ohl (Museum für Naturkunde der Humboldt-Universität Berlin, Germany).
Many thanks are also due to Stéphane Le Tirant (Montreal Insectarium, Canada) for kindly obtaining specimens
and helpful information. Edy Bhaskara (Malang, Java, Indonesia) is acknowledged for specimens, helpful informa-
tion and photos. Rob Krijns (Maastricht, Netherlands) kindly provided specimens from his cultures. The author is
indebted to sincere thanks to his good long-time friend Oskar V. Conle (Duisburg, Germany) for support in many
ways. Finally, three unknown referees are thanked for their helpful and constructive comments on the manuscript.
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