PHASMID STUDIES
volume 6, numbers 1 & 2.
June & December 1997.
Editor: P.E. Bragg.
Published by the Phasmid Study Group.
Phasmid Studies
volume 6, numbers 1 & 2.
ISSN Q966-QQ11
Contents
A brief contribution to the history of the genus Bacillus in Sicily
Simone Berni 1
An Ancient Stick Insect
Murray L. Eiland 13
The "umbrella spines" and other surface projections of some phasmid eggs and some
comments on phasmid taxonomy
John Sellick 15
Taxonomic changes relating to New Zealand stick insects
Paul D. Brock 21
A glossary of terms used to describe phasmids
P.E. Bragg 24
Reviews and Abstracts
Book Review 34
Phasmid Abstracts 35
The egg of Baculofracturn insignis (Brunner)
J.T.C. Sellick 41
A new species of Phobaeticus Brumier von Wattenwyl, from the Philippines
(Phasmatidae)
Paul D. Brock 43
Reviews and Abstracts
Book Reviews 46
Video Review 47
Phasmid Abstracts 47
Cover illustration: Male Aschiphasma anmdipes Westwood, drawing by P.E. Bragg
A brief contribution to the history of the genus Bacillus in Sicily
Simone Berni, 17 Loc. Le More di Cuna, 53014 Monteroni d’Arbia (SI), Italy.
Abstract
Some short notes on the presence of the genus Bacillus (Insecta, Phasmatodea, Bacillidae) in the Italian peninsula and
particularly in Sicily. The present paper recounts the vicissitudes that led to the discovery of new taxa in the 1980s;
provides an outline of their distribution, with notes on the relationships between the different species and a further
account of their reproduction modes. Finally, excursions to the Iblean region (south-eastern Sicily) and Marettimo
Island (Egadi Archipelago) are described, and future trips and study projects are suggested.
Key words
Bacillus, Sicily, Iblean region, Amphigonic, Hybrid, Parthenogenesis, Hybridogenesis, Gynogenesis, Androgenesis,
Species, Endemic, Extinction, Rearing, Biodiversity.
Introduction
Up to the 1960s, only one amphigonic (sexually reproducing) population of Bacillus rossius
(Rossi, 1788) in the Italian peninsula was recorded in the literature (Montalenti & Fratini,
1959) and was located in Campania. Italian researchers subsequently noted some further
findings in Liguria and Tuscany (Marina di Pisa), where both amphigonic and parthenogenetic
populations were living. With regard to B. rossius, some authors noted various sub-species
defined on electrophoretic and ootaxonomic bases; two of these are recorded in Italy (see
Figure 1). B. rossius rossius is located along the Tirrenic coasts (the coastlines of Liguria,
Tuscany, Sardinia, Campania and Lazio), and in a small area at the river Po delta, while B,
rossius redtenbacheri lives along the entire length of the Adriatic and Ionic basins and at the
tip of the Italian peninsula (Calabria, Apulia, Basilicata), including Sicily, the Eolie Islands
and a small area in southern Sardinia (see Figure 2 for a view of the Italian Districts).
Recently described taxa in Sicily
Prior to 1980 only 5. rossius and Clonopsis gallica (Charpentier, 1825) were known from
Italy and there was a considerable amount of confusion among dealers and breeders of
insects. When the efforts of researchers concentrated on southern Italy (and in particular on
Sicily) in search of bisexual stocks of B. rossius, they soon found themselves confronted with
some new species of stick insects. Within a short time period, in the Iblean region of south-
eastern Sicily, Bacillus grandii (Nascetti & Bullini, 1982), Bacillus whitei (Nascetti & Bullini,
1982) and Bacillus lynceorum (Bullini, Nascetti & Bianchi Bullini, 1984) were described.
Holotypes of new Sicilian stick insects are kept at Natural History Museum "Giacomo
Doria", Genoa, Italy. The first species immediately aroused interest because it was bisexual,
in the same manner as B. rossius. The others two species were hybrids and the interest of
the researchers was stimulated by their reproductive mechanisms, because it was clear they
were genetically free from the progenitor taxa (species which have originated the hybrids) .
Parthenogenesis, the embryonal development in the egg without the participation of the
male, was first noted in Bacillus as far back as the beginning of the 1930s, thanks to
observations made by a group of French researchers that included Cappe de Baillon, de
Vichet and Favrelle (Benazzi, 1946). These authors carried out a series of experiments by
crossing females of B. rossius, which in France is a parthenogenetic species only because
males are very rare (de Vichet, 1944), with males of the same species collected from the
Algerian coasts in bisexual populations (Favrelle & de Vichet, 1937). So-called "geographic
parthenogenesis" was presumed to take place in regions with cold climates. Later, this
assumption was contradicted by the discovery of parthenogenetic populations in areas, such
as the Iblean region, where the climate would seem to favour amphigonic populations. From
the 1960s onwards, the Italian zoologists and cytogenetists Mario Benazzi (initially) and
Phasmid Studies, 6(1): 1
Simone Bemi
Figure 1. Distribution of Bacillus rossius rossius (vertically hatched) and B. r.
redtenbacheri (horizontally hatched) along Italian coasts. After Tinti,
Mantovani & Scali (1992: 187).
Valerio Scali (currently) have completed a series of very interesting studies on the Italian and
Sicilian taxa. Their works have filled a real gap in the field of evolutionistic biology (that
branch of biology concerned with the evolution of species). They have arrived at some very
important and unexpected conclusions with respect to the "ecological niches" of some species
and the complex genetic relationships of others.
B. grandii, apart from inhabiting the Iblean region, is also found in western Sicily and
in the Egadi Archipelago, with two different sub-species situated along the coast near Trapani
and in the islands of Levanzo and Marettimo: 5. grandii benazzii and B. grandii maretimi.
In the light of these discoveries, the range of the species was further extended. It is the
Scab’s opinion (1991) that this fact opens up the possibility that B, grandii could be the
"ancestor" of the holo-Mediterranean (i.e. of the entire Mediterranean basin) forms of the
genus Bacillus, as was believed until some years ago. This "theoretical taxon" (certainly
grandiiA\k.€) must be located more towards the west of the Mediterranean basin or it may
even be extinct.
B. grandii grandii, the Iblean sub-species, is a restricted endemic and through a series
A contribution to the history of Bacillus in Sicily
Figure 2. Districts of Italy.
of attendant circumstances is itself at risk of extinction. It is located in just a few habitats
near Palazzolo Acreide, Canicattini Bagni and Floridia, in the province of Syracuse. Each
of these habitats of B. grandii grandii harbours only a small number of specimens and the
human presence represents a continuous threat, especially considering the local custom of
burning or drastically trimming hedges, the only refuge for phasmids. The second major
aspect creating the risk of extinction in B. grandii grandii is the incessant competition with
Phasmid Studies, 6 ( 1 ): 3
Simone Bemi
the sympatric (two or more species which live together) B. whitei with which it mates
regularly, as confirmed by Scali, Mantovani and Tinti (1991: 104). This fact causes a
substantial waste of the insects’ reproductive potential, only partly balanced by the sex ratio
in favour of males.
B. whitei, a hybrid of B. rossius redtenbacheri and B. grandii grandii, has lately been
the subject of some interesting smdies that have thrown light on its genetic characteristics.
Such characteristics mean that B. whitei is a taxon that is unique in the whole of the animal
kingdom. By means of a series of crosses with the three sub-species of B. grandii,
researchers have found the contemporaneous presence of parthenogenesis, hybridogenesis and
gynogenesis, and have also reported occasional cases of androgenesis (Scali, Mantovani &
Figure 3. Reproduction modes in the genus Bacillus', after Tinti in Veroli (1995: 26).
Phasmid Studies, 6 ( 1 ): 4
A contribution to the history of Bacillus in Sicily
Tinti, 1991) (see Table 1 and Figure 3 for a general scheme of reproduction modes in the
genus Bacillus).
This is the first practical confirmation of the evolutionistic theory put forward by some
authors on hybridogenetic organisms. For this reason, this representative of Iblean fauna
takes on enormous importance, as it testifies to the genuine possibility that an amphigonic
organism, evolved to the hybridogenetic stage, might select parthenogenesis as a further
adaptation. This is also important because, for the first time, hybridogenesis is being
considered as a vital genetic-evolutionistic adaptation. This could have highly interesting
reverberations for other animal classes and call for renewed discussion of the very idea of
"species" (Veroli, 1995).
B. lynceorum, the longest species in the genus, is the triploid rossius-atticus-grandii,
probably a cross between B. grandii grandii and a diploid FI hybrid B. atticus x B. rossius
redtenbacheri which (unusually) produced fertile eggs (Brock, 1991: 20; Manaresi et al.,
1992).
In the 1980s, together with the discovery of new Sicilian taxa, some further locations
in Italy for Bacillus atticus Brunner von Wattenwyl, 1882 were discovered, which was
previously only recorded from Attic, Epirus and Peloponnese (in Greece) according to
Nascetti and Bullini (1982), La Greca (1984, 1996a), Agostini and Scali (1989) and on the
Dalmatian coasts (Muller, 1957). During the Pleistocene era (La Greca, 1996a), this taxon
arrived in the Iblean region by crossing what is now Apulia and Calabria and spread
throughout the whole of Sicily. In fact, the micro-plate of African origin that formed the
Iblean region, during the Pliocene era, was again separated from the north of Sicily (see
Figure 4). B. atticus on the Italian coasts displays a genetic differentiation from its
counterpart in Greece and in the Dalmatian area. In 1982, the researchers Nascetti and
Bullini named the Italian population as new sub-species B. atticus caprai. Strangely, this
subspecies is not cited in Failla et al. (1994).
1
Species
Mode of reproduction
Bacillus rossius
amphigony (*), parthenogenesis (**)
Bacillus atticus
parthenogenesis
Bacillus grandii (***)
amphigony
Bacillus whitei
parthenogenesis, hybridogenesis, gynogenesis androgenesis
Bacillus lynceorum
parthenogenesis
Bacillus rossius-grandii grandii
hybridogenesis, androgenesis
Bacillus rossius-grandii benazzii
hybridogenesis, androgenesis, gynogenesis
Table 1. Reproduction modes in the Italian representatives of the genus Bacillus.
(*) Central and southern Italy; (**) Central and northern Italy; (***) All three
subspecies of the taxon.
Of additional interest were the fmdings of specimens of the natural hybrid Bacillus
atticus-rossius, along the coast near Alimini, in Apulia. However, these insects are
completely sterile. Again, with respect to the Sicilian phasmids, it is doubtful whether the two
inter-species hybrids B. rossius- grandii grandii of the Iblean region and B. rossius- grandii
Phasmid Studies. 6(1): 5
benazzii of north-west Sicily should really be considered species in every respect. In
accordance with the interpretation given in the recent "Checklist delle specie della fauna
italiana" (Failla etal, 1994: 19-20) - probably the most authoritative reference among those
cited - authors are inclined to assign the status of species, at least temporarily (these hybrids
are in course of study), although these populations seem to be unable to maintain themselves
without continuous crosses with the progenitor taxa. In fact in zoology a species is the basic
unit: an individual entity of a population, the members of which show the same chromosome
complement; they are able to reproduce themselves, giving origin to fertile progeny; such
progeny must be able to maintain themselves. (See Figure 5 for a general view of the
Sicilian taxa’s distribution).
Figure 4.
What is now Italy, as it was in the Pliocene era; after Blanc in La Greca
(1996a: 24).
Notes on two preliminary excursions to Sicily - October 1996
In October 1996, in collaboration with the Ente Fauna Siciliana, some adult specimens of B.
whitei (see Figure 6) were found, at night, near Isola Coco, alongside the old Syracuse-
Vizzini-Ragusa railway, on some bramble plants (Rubus spp.). The plants were in the
undergrowth, near the river Anapo. Although very common in the Iblean region, B. whitei
Phasmid Studies, 6 ( 1 ): 6
A contribution to the history of Bacillus in Sicily
had never been recorded as inhabiting this area. The only record in the scientific literature
for the Anapo Valley seems to have been in Bullini and Nascetti (1987), but this is for the
genus Clonopsis. Some specimens of B. whitei, collected for study, are at present being kept
in captivity at room temperature and under natural photoperiod. Attempts are being made
to understand its capacities for adaptation and longevity in captivity. With respect to
foodplants for rearing, Scali (1991: 403) suggests that a mixed diet of bramble and lentisk
could be the best solution for B. grandii. Other specimens of B. whitei were collected in a
bramble-bush in the suburbs of Palazzolo Acreide (province of Syracuse). During a further
excursion along the road between Canicattini Bagni (province of Syracuse) and the cross-road
for Palazzolo Acreide, some specimens of B. lynceorum were collected. This species is easily
recognizable thanks to its remarkable size (up to 105mm in length), the clear granulation of
its meso and metathorax, its stocky cerci and the typical notch at the distal end of its
subgenital plate. B, lynceorum is also exclusively endemic to the Iblean region.
Figure 5. Distribution of Bacillus spp. in Sicily; after Montovani & Scali, as modified
by La Grece (1996a: 32).
The trip to the Iblean region came to an end in western Sicily along the Trapani coast,
between the localities of San Vito Lo Capo and Scopello where B. grandii benazzii were
noted and in Marettimo (Egadi Islands) where B. grandii maretimi (see Figure 7), is found.
During an excursion in Marettimo several green nymphs (at first and second instar) of B.
grandii maretimi were found on lentisk. These two sub-species, in fact, feed only on lentisk
(Pistacia lentiscus Linnaeus), in contrast with B. grandii grandii of the Iblean mountains
which feeds principally on bramble (Rubus spp.) or plants of Rosa spp.
Studies and prospects
With regard to the Iblean region (see figure 8), the author plans for 1997 foresee a more
detailed study along the Anapo and Cassibile rivers in search for new habitats of B. whitei,
Phasmid Studies, 6 ( 1 ): 7
Figure 6. Female Bacillus whitei on bramble.
Phasmid Studies, 6(1): 8
A contribution to the history of Bacillus in Sicily
B, lynceorum and the hybrid B. rossius-grandii grandii. It is very difficult to recognize this
hybrid because it is morphologically similar to Bacillus whitei (Failla et al. , 1994: 22). This
taxon, doubted by some authors, is different from B. whitei in being hybridogenetic as
opposed to parthenogenetic. Consequently, while B. whitei is completely free of males of B.
grandii grandii for purposes of reproduction, B, rossius-grandii grandii requires continuous
crosses with "progenitors".
Figure 7. Bacillus grandii maretimi, three nymphs on lentisk.
A second - and no less important - purpose concerns the conservation of the taxon B.
grandii grandii. As previously stated, this species is in serious risk of extinction. We may
assume that the best method of conservation for the taxon is the protection of its environment:
hedges of bramble bushes. Unfortunately, the extensive use of fire in order to remove
unwanted hedges in the Iblean territory is a cause of death for this insect. If these natural
barriers were to be destroyed manually, the insect would at least have the possibility of
escape during the night.
Phasmid Studies, 6 ( 1 ): 9
Simone Bemi
The hedge problem has already been discussed in relation to interventions aimed at
protecting natural environments, with WWF Italy playing a major role. During the early
1990s, several thousand shrubs of Prunus, Myrtus, Rubus, Cistus, Pyrus, Crataegus, Rosa,
Pistacia, Erica and other characteristic plants of the Mediterranean environment, were planted
across Italy. Thus, efforts are being made to revive at least a part of a territory that has been
sorely impoverished in earlier decades (Francescato, 1989).
Figure 8. The Iblean area of Sicily.
Biodiversity in a natural environment is the result of a long developmental process. The
most precious aspect of the resources in a natural environment is provided by the endemic
species, that is, by the living creamres geographically localized and currently at risk of
Phasmid Studies, 6 ( 1 ): 10
A contribution to the history of Bacillus in Sicily
extinction. It should be remembered that an incidental loss of these species would be an
incalculable loss for science and utterly irredeemable (La Greca, 1993, 1996b). The situation
in Marettimo is more stable, in contrast with the Iblean region; B. grandii maretimi is present
in a large numbers and there are currently no obstacles to its survival.
Acknowledgements
I am grateful to Lorenzo Aiello for drawing figures 1 and 3-7. Thanks to: Giovanni Aliotti,
Phil Bragg, Paul Brock, Marcello La Greca, Carmelo Milluzzo, Leonardo Parisi, Roberto
Poggi, Bruno and Franco Ragonese, Ivor Rowan, Paolino Uccello; for support and valuable
suggestions.
References
(*) The bibliographic notes prefixed by an asterisk denote works not examined by the author;
they have been cited by other authors.
Agostini, N. & Scali, V. (1989) Segnalazione di nuove stazioni italiane di Bacillus atticus Brunner, 1882: Isole
Tremiti, Gargano e Sicilia. Bollettino Societa entomologica italiana, Genova, 121(1): 10-12.
Benazzi, M. (1946) Qualche osservazione sul fasmide Clonopsis gallica Charp. Atti della Sezione Agraria, Siena.
Reale Accademia dei Fisiocritici, 10: 20-25.
Brock, P.D. (1991) Stick insects of Britain, Europe and the Mediterranean. Fitzgerald Publishing, London.
(*)Bullini, L. & Nascetti, G. (1987) Genetic and taxonomic smdies on the genus Clonopsis with the description of
a new species (Phasmatodea, Bacillidae), Bollettino Istituto Entomologia Umversitd, Bologna, 4: 325-353 [cited in
Brock, 1991].
Failla, M.C., La Greca, M., Lombardo, F., Messina, A., Scali, V., Stefani, R., & Vigna Taglianti, A, (1994)
Blattaria, Mantodea, Isoptera, Orthoptera, Phasmatodea, Dermaptera, Embioptera. In\ Minelli, A., Ruffo, S. & La
Posta, S. (eds) Checklist delle specie della fauna italiana, 37. Bologna, Calderini.
(*)Favrelle, M. & Vichet, G. de (1937) Resultats de la fecondations, par un male d’Algerie, de femelles
parth6nog6n6tiques ffangaises du Bacillus rossii (Phasmidae). Compte Rendu Academie des Sciences, 204: 1899-1900
[cited in Benazzi, 1946, and in others].
Francescato, G. (1989) Facciamo siepe contro I’avanzata dei veleni. Natura Oggi, 7(1): 54-61.
(*)La Greca, M. (1984) L’origine della fauna italiana. Le Scienze, 187: 66-79 [cited in Agostini & Scali, 1989].
La Greca, M. (1991) Le molte facce della Politica Ecologica Italiana e la tutela degli ambienti naturali. Natura e
Montagna, 38(1-2).
La Greca, M. (1993) I problem! della conservazione della fauna e la politica gestionale degli ambienti naturali. Atti
e Memorie delVEnte Fauna Siciliana, 1: 35-42.
La Greca, M. (1996a) Origine della fauna iblea. In: Ragonese Bruno (ed) La fauna degli iblei, atti del convegno su
la fauna degli iblei, 13-14 maggio 1995, Syracuse, Ente Fauna Siciliana.
La Greca, M. (1996b) Gli endemismi: un prezioso bene naturale e culturale da tutelare. Grifone, 5: 1-2.
(*)Manaresi, S., Marescalchi, O. & Scali, V, (1992) The chromosome complement of the hybrid Bacillus whitei
complex (Insecta, Phasmatodea). The paleo- and neo- standard karyotypes - The repatterned cytotypes. Cytologia,
57: 101-109, 111-119 [cited in Veroli, 1995].
(*)Montalenti, G. & Fratini, L. (1959) Observations on the spermatogenesis of Bacillus rossius (Phasmoidea), XV
International Congress of Zoology Proceedings, London, 749-750 [cited in Scali, 1964, 1970].
(•)MuUer, G. (1957) Faunisticka istrazivanja sjeverodalmatinskih otoka Dugi otok i Kornati (1925-1927).
Orthopteroidea, Coleoptera i Formicidae - Jugol. Akad. Znanosti i Umjemosti, Odjel prirodne nauke. Acta Biologia,
1: 187-218 [cited in Agostini & Scali, 1989].
(*)Nascetti, G. & Bullini, L. (1982) A new phasmid from Italy: Bacillus atticus caprai (n. subsp.) (Cheleutoptera,
Bacillidae). Fragmenta Entomologica. 16: 143-159 [cited in Agostini & Scali, 1989].
(*)Nascetti, G. & Bullini, L. (1983) Differenziamento genetico e speciazione in fasmidi dei generi Bacillus e
Clonopsis (Cheleutoptera, Bacillidae). Atti XII Congresso Nazionale Italiano di Entomologia, 2: 215-223 [cited in
Agostini & Scali, 1989].
Scali, V. (1964) Modalita riproduttive della popolazione di Bacillus rossius (Rossi) dei dintorni di Pisa. Rendiconti
Scienze fisiche, matematiche e naturali, Accademia dei Lincei, 36; 311-314.
Scali, V. (1968) Biologia riproduttiva del Bacillus rossius (Rossi) nei dintorni di Pisa con particolare riferimento
all’influenza del fotoperiodo. Atti Societa Toscana di Scienze Naturali, 75: 108-139.
Phasmid Studies, 6(1): 11
Simone Bemi
Scali, V. (1970) Obligatory parthenogenesis in the stick insect Bacillus rossius (Rossi). Rendiconti Scienze fisiche,
matematiche e naturali, Accademia dei Lincei, 49: 307-314.
Scali, V. (1991) Un nuovo insetto stecco (Phasmatodea) della Sicilia: Bacillus grandii benazzii (n. subsp.). Frustula
entomologica, 12: 397-408.
Scali, V., Mantovani, B. & Tinti, F. (1991) Primi dati sull’ibridogenesi, androgenesi e ginogenesi di Bacillus whitei
Nascetti & Bullini (Insecta Phasmatodea). Frustula entomologica, 12: 103-108.
Tinti, F., Mantovani, B. & Scali, V. (1992) Caratterizzazione allozimatica di popolazioni di Bacillus rossius
deiritalia centro-meridionale e della Sicilia. Bollettino Societa Entomologica Italiana, 123(3): 184-194.
Veroli, R. (1995) Gli insetti stecco, un caso emblematico della tassonomia. La ricerca di Valerio Scali e dei suoi
collaboratori, L’Aquila-Roma, Japadre.
(*)Vichet, G. de (1944) D6couverte dans le sud de la France d’une station de miles de Bacillus rossii Fabr. Bulletin
Societe Linn. , 13: 92-94 [cited in Scali, 1968].
Phasmid Studies, 6(1): 12
An Ancient Stick Insect
Murray L. Eiland, mo Eagle Nest Court, Danville, CA, 94506, USA.
Abstract
This article proposes that a Chinese tree root carving of the fourth to third century BC is based on a stick insect.
Key words
Phasmida, Chinese, Antiquities.
This artifact (Figure 1) is noted today as being the oldest surviving root carving from Ancient
China. The head and body of the animal is made of smoothed tree root, while the legs are
rendered in bamboo, which clearly show their sectional structure. The entire creature is
lacquered maroon red. The head is certainly feline, with large orange (ground) and red
(pupil) eyes. Its teeth, in a formidable fence-like array, may be taken in an almost comical
manner. The rest of the animal is anything but threatening. The right front leg bears a snake
that wriggles up the length of the limb, while on the right hind leg a short snake bites a
tailless lizard. The left front leg bears a lizard clutching the head of a (stylized) bird in its
jaws, which appears on this drawing, on such a limited scale, as a floral device. The top of
the left hind leg bears a small but easily identified cicada.
Figure 1. Tree root carving with bamboo and lacquer, height 32cm; length 69.5cm. Eastern
Zhou, Warring States Period (4th-3rd century BC), Chu state. Excavated from tomb Ml at
Mashan in Jiangling county, Hubei province, and now in the Jingzhou Regional Museum,
Hubei province (Beijing, 1985: pi. 43:1; Goepper, 1995: no. 75).
While figures such as dragons, which have a long history in Chinese art, can be
interpreted using written sources and parallels with later folklore, the nature and identity of
this figure remains elusive, as do the figures that adorn its body. Were these figures common
in Chu culture? When dealing with material from this period little can be taken as certain.
Not only is there a dearth of textual material to aid in interpretation, but the namre of root
Phasmid Studies, d(l): 13
Murray L, Eiland
carvings as a group is also obscure, as very few have survived. From its position in the tomb
where it was found, above the head of the female occupant, this carving could be seen as a
kind of guardian, to "ward off evil" (Rawson, 1996: 143). At this period in Chinese history
it is clear that there was a growing interest in "shamanism", and graves contained offerings
other than bronze. At the same time the art of the Eastern Zhou is unusually rich, with a
variety of motifs and geometric devices. As a result, many pieces were clearly meant to be
appreciated on many levels (as this root carving contains a number of figures that are related
to a greater or lesser degree). At the same time, using modem art-historical taste as a guide
(which, it may be noted, is often quite different from the aesthetics of antiquity), this figure
could be the favourite of the owner that shows no more than the mind-set of the occupant.
This creature has been interpreted by modem art historians as an "imaginary beast",
although there is compelling evidence to suspect that it is a rendition of a stick insect, a
creature that, to many museum curators, would indeed seem imaginary. While there are a
number of particular features of this figure that indicate it is composed from a variety of
animals, from the feline head to the smaller animals on the legs (it should be noted that there
are four legs rather than six in insects), the overall morphology of this animal is clear. It has
a very long thin body with long thin legs. In the case of the latter it is interesting to note that
one leg is clearly projected forward into space, which is consistent with common behaviour
of many phasmid species (as noted below).
Tlie treatment of the head of the animal also deserves special consideration. While it
is clear that it is based upon a feline model, one should not then quickly assume that this
factors against an insect-inspired origin for the animal as a whole. It should be noted that
many phasmid species have a combination of mouthparts and eyes that lead many who are
unfamiliar with the habits of this order to conclude that phasmids are capable of inflicting
considerable damage, and that, like mantids, they are carnivorous. As in many art objects
of antiquity - and today - attention to detail is often sacrificed to capture the perceived
qualities of the object of interest. Instead of depicting large compound eyes and complex
mouthparts, the artist may have chosen to simplify his subject and portray a more easily
conceived form.
It is from this latter point that one may pose the question, why is this root carving not
of a mantis? While it is clear that the Chinese - along with many other cultures - have long
venerated mantids as animals endowed with special properties due to their unique form, this
carving lacks the main attribute of a praying mantis, the raptorial legs. While not venerated
in modem societies today, phasmids show many features that may have inspired intrigue.
Many phasmids exhibit an immobility reflex when threatened, where the animal projects its
limbs forward and remains motionless for periods of time. Many species also have a swaying
pattern of motion, while others stop and waft their legs in the air for anchorage if they are
placed upon a surface with little opportunity for vertical movement. Also of interest are the
many patterns of escape which different species display, from moving backwards to dropping
and flying. Is any of this behaviour recorded in the positioning of this root carving? Without
further information, this aspect may remain conjectural. However, it is with such behaviour
in mind that one can appreciate why a phasmid would be portrayed in antiquity. Once we
understand that this creature is a stick-insect, we can also understand why two snakes, two
lizards, a bird, and a cicada are associated with this animal. These are all animals that one
would expect to find in a phasmid ’s domain.
References
Beijing (1985) Jiangling Mashan Chu mu. Beijing.
Goepper, R.(1995) Das Alte China: Menschen and Goner im Reich der Mine. Munich.
Rawson, J. (1996) Mysteries of Ancient China: New Discoveries from the Early Dynasties. British Museum Press,
London.
Phasmid Studies, 6(1): 14
The "umbrella spines" and other surface projections of some phasmid eggs
and some comments on phasmid taxonomy
John Sellick, 31 Regent Street, Kettering, Northants, NN16 8QG, U,K.
Abstract
Short flat-topped spines are found in four tribes of Phasmida, widely separated in the traditional subdivision of the
order. Pinnulae occur in Phy Ilium siccifolium. A survey is given of other surface projections of various types found
in different families. A table of subfamilies is given, indicating some taxonomic problems.
Key words
Phasmida, egg structure, spines, hairs, pinnulae. Bacteria, Baculum, Creoxylus, Datames, Dinopkasma, Epidares,
Haaniella, Hoploclonia, Paraphasma, Phenacephorus, Phyllium, Pseudophasma, Sipyloidea, Stratocles.
Umbrella spines
I first noticed these structures when preparing s.e.m. images of two species of Baculum for
my 1980 thesis. They stand around 0.15mm from the general egg capsule surface with a
relatively narrow stalk and then expand into a more or less complex flattish top, the rim of
which is composed of fused roughly spherical units. Reading recently Frederic Langlois’
account (1995) of the egg of Stratocles tessulata (Olivier) [as Stratocles variegatus (Stoll)]
I saw there the same structures in an egg of a distinctly different group of phasmids.
Langlois drew attention to the similarity of these structures in the two genera. He described
them as "structures en forme de parapluie" (umbrella-like structures). Figure 1 shows the
s.e.m. form of these "umbrellas".
I had noted their occurrence in Paraphasma rufipes (Redtenbacher), which like
Stratocles is in the tribe Stratocleini, and in Pseudophasma bispinosa (Redtenbacher) in the
Pseudophasmatini. Going to my reference collection I found them also in Baculum cuniculum
(Westwood), Baculum insignis (Wood-Mason), Baculum insueta (Brunner), Baculum PSG 144
and Baculum PSG 157 in the Baculini, all of which are Baculum (iii), the third of the four
strikingly different forms of eggs found within this "genus" (Sellick 1997). They are also in
the Creoxylus sp. supplied to me by Oliver Zompro (his no. 86) which is in the Xerosomatini.
Here they are confmed to the raised lines on the capsule surface, and form a ring on the
operculum. I have not been able to detect them in all the species in my collection of Baculum
(iii) or of Creoxylus. Stratocleini, Pseudophasmatini and Xerosomatini are closely related
tribes (subfamily Bacunculinae), but in the traditional classification the Baculini are far
separated from them (subfamily Phasmatinae). The first two tribes are Areolatae, whilst the
Baculini is in the Anareolatae. Where else are these to be found? Similar structures occur
in Phyllium celebicum (de Haan). These have shorter stalks (about 0.07mm) and complex
heads (fig. 2A). The curious crusty surface of the egg of Bacteria PSG 152 (Subfamily
Bacteriinae) is composed of the tessellated top plates of large umbrella spines (fig. 3). Like
the others they stand around 0. 15mm above the surface, their top plates being up to 0.35mm
across, compared with 0.15- 0.25mm in those species in other tribes where they do not form
a full surface layer. The top plates are particularly large in a single layer surrounding the
micropylar plate and along the opercular collar, where they form a series of open scales.
Whilst the spines of Stratocleini, Xerosomatini and Baculini are so similar that they seem to
indicate a close relationship, it is possible that those of Bacteriinae and Phylliidae with
different dimensions are an accidental similarity of different origin.
The surface of the egg of Phenacephorus comucervi (Brunner) (Lonchodinae) appears
to be a pattern of rings with central depressions. Sectioning shows that these rings are
produced by surface extensions with some similarity to umbrella spines in that they have
stalks and an expanded head. They surround a thickening of the translucent compact layer
of the capsule (fig. 5B). All the heads are however united into the ring formations, leaving
Phasmid Studies, 6(1): 15
J. Sellick
irregular openings which mark the circumferences of the rings (fig. 5A).
Figure 1. Umbrella spines.
A-D Side views: A. Baculum thaii. B. Baculum cuniculum.
variegatus.
E-H Surface views of B. thaii.
C & D. Stratocles
All traced from s.e.m. images (A-B, E-H Sellick 1980; C-D Langlois 1995).
Figure 2. Phy Ilium.
A. Flat-headed spines in P. celebicum showing variety in shape and size of heads.
B & C. Pinnulae of P. siccifolium: B. side view of projecting portion. C. view
showing base.
Pinnulae
I used this term in Sellick 1978 to describe the feather-like extension of the capsule and
Phasmid Studies, 6 ( 1 ); 16
Umbrella spines and other surface projections on phasmid eggs
operculum surface characteristic of Phy Ilium siccifolium (Linnaeus), but not so far found in
any other species. These (figs. 2B & 2C) originate in a base some 0.5mm long which runs
along the surface of the egg tapering from a broad origin almost to a point; it then extends
outwards as a one- or two-branched flattened structure up to 1.6mm long and around 0.5mm
wide which curves somewhat back in the direction of the base.
Figure 3. Umbrella spines forming the surface of the egg of Bacteria PSG 152.
A. Section through the micropylar plate.
B. Surface view of the area at the front of the micropylar plate.
C. Scale-like plates of the opercular collar seen in transparency, stalks of the plates are
shaded.
D. Section of this region.
Drawings using a light microscope, pig = sunken pigmented surface layer over the
plate, imp = internal micropylar plate, off = opercular fringe.
A B
Figure 4.
A. Formation of collar fringes in Diesbachia,
B & C. Hairs of Datamini. B. Epidares. C. Datames.
Phasmid Studies, 6 ( 1 ): 17
J. Sellick
2
❖
J"
A
B
Figure 5. The surface of Phenacephorus comucervi.
A. Surface view showing "pit" and "ring".
B. section through a pit and ring system.
Fringes of the capsule collar
A number of genera of the Necrosciidae possess fringes on the collar which surrounds the
operculum. These are of various lengths, ranging from 0.3mm in Orxines macklottii (de
Haan), through 0.5mm in Acacus sarawacus (Westwood) and 0.6mm in Diesbachia tamyris
(Westwood), to 0.65mm in Centrophasma hadrillus (Westwood). The fringe material is
traversed by some sixty thickened struts, each about 0.2mm wide and it then splits into
separate fringes, each fringe containing usually two or three of these struts (fig. 4A). These
are all true capsule outgrowths, and must not be confused with the fringe of Trachythorax
maculicollis (Westwood), which is produced by a splitting away of the surface capsule layer.
What appears to be a collar fringe in various species of Baculum, particularly PSG 157, is
produced by variously developed umbrella spines and is opercular and not capsular.
Egg capsule hairs
Hairs of various types occur sporadically throughout the groups of phasmids. In Orobia sp.
(from Madagascar) [Pygirhynchinae] there are short fine hairs ca. 0.03mm long. Three
species of Haaniella [Heteropteryginae, Heteropterygini] show fine hairs of different lengths:
0.02mm in H. echinata (Redtenbacher), 0.07 mm in H. grayii (Westwood) and 0,13mm in
Zompro’s no. 30. Similar fine hairs, 0.08mm long, are found in Hoploclonia gecko
(Westwood) [Heteropteryginae, Obrimini]. Longer hairs, 0.18mm long, are found over the
capsule and operculum surface of Sipyloidea ?sipylus PSG 4. In the Heteropteryginae,
Datamini three species show forms of hooked hairs. In Datames oileus (Westwood) these are
sparse and 0.22mm long, expanded at the tip to form a three- (possibly four-) pronged top
(fig. 4C). Epidares nolimetangere (de Haan) has a dense covering of long (0.4 - 0.45mm)
hairs each with a single hook at the tip (fig. 4B). Dinophasma guttigera (Westwood)
[Aschiphasmatinae, Aschiphasmatini] has a dense fine covering of 0.2mm hairs.
Phasmid Studies, 6 ( 1 ): 18
Umbrella spines and other surface projections on phasmid eggs
Comments on phasmid taxonomy
The division of the Phasmida into two on the basis of the presence or absence of triangular
areas (the "area apicalis") on the mid- and hind-tibiae has been brought into question by a
number of authors (e.g. Kristensen 1975, Key 1974, Roberts 1974), although retained by
Kevan (1982) who, after the removal of Timema as the suborder Timematodea, divided his
other suborder, the Phasmatodea, on conventional lines. Kristensen merely criticised the
value of the area apicalis as a taxonomic criterion, pointing out that it may be sclerotised or
not, elevated or depressed, and delimited by carinae, rows of spines, or have neither of these.
He made no suggestion for regrouping within the order. Key suggested that the criterion for
membership of the "Areolatae" should be the presence of a male vomer, on which basis he
would transfer the Necrosciidae into that suborder. Roberts found vomers in Lonchodinae
and Bacteriinae, both traditionally Anareolatae. These latter three groups are themselves at
present in the two different superfamilies of the Anareolatae.
As an illustration of the chaos of phasmid classification, table 1 shows the 23
subfamilies of the Kevan classification, with some of their characteristics. Some of the names
have been corrected following Bragg (1997), as they also have been in the above account.
In no case is it to be taken that any of these characteristics is found throughout the subfamily
in question.
Infraorder
Superfamily
Family
Subfamily
1
2
3
4
5
Timematodea Timematoidea
Timematidae
—
+
-
-
X
ab
Bacillidea
Bacilloidea
Bacillidae
Bacillinae
+
+
-
0/C
+
Pygirhynchinae
+
+
-
c
ab
Heteropteryginae
+
+
-
0
+/ab
Pseudophasmatidae Aschiphasmatinae
+
+
-
c
ab
Korinninae
+
+
-
0
+
Pseudophasmatinae
+
+
+
0
+/ab
Phyllioidea
Phylliidae
—
+
+
+
c
ab
Phasmatidea
Heteronemioidea
Necrosciidae
—
ab
+
-
0/C
+/ab
Heteronemiidae
Heteronemiinae
ab
-
-
0/C
+
Libethrinae
ab
-
-
0
+
Lonchodidae
Lonchodinae
ab
+
+?
0
+
Menexeninae
ab
-
-
0/C
+ /ab
Pachymorphidae
Gratidiinae
ab
-
-
0/C
+/ab
Pachymorphinae
ab
-
-
c
ab
Palophidae
—
ab
-
-
0
ab
Phasmatoidea
Bacteriidae
Cladoxerinae
ab
-
-
-
-
Bacteriinae
ab
+
+
0
+/ab
Phasmatidae
Platycraninae
ab
-
-
0/C
+ /ab
Xeroderinae
ab
-
-
-
-
Eurycanthinae
ab
-
-
0
+
Tropidoderinae
ab
-
-
c
ab
Phasmatinae
ab
-
+
0/C
+/ab
Table 1. Phasmid subfamilies.
1 Area apicalis on tibiae
2 Male vomer
3 Egg umbrella spines
4 Internal micropylar plate type; X = no gap; O = open gap; C = closed gap
5 Median line: + = present in at least some genera; - = presence not known; ab = definitely absent in genera
examined.
Phasmid Studies, 6(1): 19
7 . Sellick
References
Bragg, P.E. (1997) Phasmids of Borneo, Ph.D. thesis, University of Nottinghanoi.
Kevan, D.K.McE. (1982) Phasmoptera, in Parker, S.F. (Ed.) Synopsis and Classification of Living Organisms,
Vol.2, McCjraw Hill, New York et al. 379-382.
Key, K.H.L. (1974) in C.S.I.R.O. Insects of Australia, Supplement, Melbourne University Press.
Kristensen, N.P. (1975) The phylogeny of the hexapod ’orders’. A critical review of recent accounts. Zeitschrift fur
die Zoologische Systematik und Evolutionsforschung, 13: 1-44.
Langlois, F. (1995) L’oeuf de Stratocles variegatus (Stoll, 1813). Le Monde des Phasmes, 29: 3-9.
Roberts, H.R. (1974) Footnote in Key 1974.
Sellick, J.T.C. (1978) The eggs of leaf insects (Insecta: Phasmida). Zoological Journal of the Linnean Society, 63:
249-258.
Sellick, J.T.C. (1980) A study of the eggs of the insect order Phasmida with panicular reference to the taxonomic
value of egg structure in this group, Ph.D. thesis, University of London, 349 pp.
Sellick, J.T.C. (1997 - in press) Descriptive terminology of the phasmid egg capsule, with an extended key to the
phasmid genera based on egg structure. Systematic Entomology, 22: 101-126.
Phasmid Studies, 6(1): 20
Taxonomic changes relating to New Zealand stick insects
Paul D. Brock, "Papillon", 40 Thorndike Road, Slough, SL2 ISR, U.K.
Abstract
Following research on New Zealand stick-insect type specimens four new synonyms have been identified, as follows:
Pachymorpha finitima Brunner von Wattenwyl, 1907 and Tectarchus diversus Salmon, 1954 are synonyms of
Tectarchus huttoni (Brunner von Wattenwyl, 1907) (which has been transferred from the genus Pachymorpha Gray).
Pachymorpha bouvieri Brunner von Wattenwyl, 1907 is a synonym of Mimarchus annulatus (Hutton, 1898); and
Clitarchus interruptelineatus Brunner von Wattenwyl, 1907 is a synonym of Clitarchus hookeri (White, 1846).
Key words
New Zealand, Salmon, Brunner von Wattenwyl, Naturhistorisches Museum Wien, synonym, Phasmida.
Introduction
Whilst revisiting the Naturhistorisches Museum Wien (NHMW) in 1996, I took the
opportunity of examining Brunner von Wattenwyl’ s New Zealand taxa unfortunately
overlooked in Salmon’s excellent book (1991). I had already seen type material in Paris
(MNHN) and believed that the Pachymorpha species described by Brunner von Wattenwyl
in 1907 had been incorrectly placed in that genus.
My research has made certain taxonomic changes necessary, as listed in the following
section. Listings of species include only the most important references or a change of genus
made by a later author. Museum codens are also given below:
BMNH The Natural History Museum, London, England.
CMNZ Canterbury Museum, Christchurch, New Zealand.
MNHN Museum National d’Histoire Naturelle, Paris, France.
MONZ Museum of New Zealand, Wellington, New Zealand.
NHMW Naturhistorisches Museum Wien, Wien, Austria.
ZMUH Zoologisches Instimt und Zoologisches Museum, Universitat Hamburg, Hamburg, Germany.
Tectarchus Salmon, 1954
Tectarchus Salmon, 1954: 161. Type species: Tectarchus diversus Salmon, 1954: 163,
designated by Salmon, 1954: 162 [= Tectarchus huttoni (Brunner von Wattenwyl), 1907:
213; see below].
Tectarchus huttoni (Brunner von Wattenwyl) comb.n.
Pachymorpha huttoni Brunner von Wattenwyl, 1907: 213. Syntype series: 6, 299, New
Zealand (NHMW, No. 383); c3, 9, New Zealand: Nelson (MNHN).
Pachymorpha finitima Brunner von Wattenwyl, 1907: 215. Holotype 9, New Zealand,
[18J48-52, leg. Petit, "Mus. Paris” (NHMW, No. 380). syn.n. [No types of this
species were found in MNHN] .
Tectarchus diversus SdXmon, 1954: 163, pi. 7: 1-2, pi. 8: 1, 2,4,8, pi. 9: 1 &5. Holotype
& paratypes 66 & 99, New Zealand (several localities) (MONZ, including coll.
Salmon; also in coll. G. Ramsey), syn.n.
Mimarchus Carl, 1913
Mimarchus Carl, 1913: 22. Type species: Mimarchus tarsatus Carl, 1913: 23, by monotypy.
Mimarchus annulatus (Hutton)
Pachymorpha annulata Hutton, 1898: 162. Holotype 9, New Zealand: Dunedin (CMNZ).
Mimarchus annulatus Hutton; Salmon, 1991: 96.
Phasmid Studies, 6(1): 21
P.D. Brock
Pachymorpha bouvieri Brunner von Wattenwyl, 1907: 214. Syntype series: 366, 399,
New Zealand: Invercargill, leg. Burr; 6, 29 9, New Zealand: Nelson, 1876, leg.
Filhol, "Mus. Paris"; 6, no locality (NHMW, No. 378); 66 & 99, New Zealand
(MNHN). syn.n.
Three further females in NHMW, without identification labels, are not regarded as part of
the type series.
Clitarchus StM, 1875
Clitarchus Stal, 1875: 34, 82. Type species Clitarchus laeviusculus Stal, 1875: 82, by
designation of Kirby, 1904: 339 [= Clitarchus hookeri (White), 1846].
Clitarchus hookeri (White)
Pkasma hookeri White, 1846: 24, pi. 6: 6. Holotype 9, New Zealand (BMNH).
Bacillus hookeri (White); Westwood, 1859: 14.
Clitarchus hookeri (White); Stal, 1875: 83.
Clitarchus laeviusculus Stal, 1875: 82. Syntype series: 49 9, New Zealand, leg. Boucard
(NHMW, No. 443). (synonymised by Ragge, 1965: 39).
Bacillus coloreus CoXemo, 1885: 151. 9, New Zealand: Pourerere, near Blackhead, 1884,
leg. W. Scott, (synonymised by Brunner von Wattenwyl, 1907; 237).
Bacillus minimus Colenso, 1885: 153. New Zealand: Norsewood. (synonymised by
Salmon, 1991: 82).
Clitarchus reductus Hutton, 1899: 55. 9, New Zealand: Canterbury (CMNZ).
(synonymised by Salmon, 1991: 82).
Clitarchus interruptelineatus Brunner von Wattenwyl, 1907: 236, pi. 10: 4a-d. Syntype
series; d, 29 9, New Zealand (NHMW, No. 445); 6, New Zealand: Great Barrier
Island (ZMUH). syn.n.
Summary
Apart from noting new synonyms, there is only one significant change to make to Salmon’s
book, i.e. references to Tectarchus diversus should read Tectarchus huttoni (for main section
on this species, see pages 101-105 of Salmon, 1991).
Whilst a number of species described by Brunner von Wattenwyl (1907) and
Redtenbacher (1906; 1908) have been synonymised, I consider that once detailed studies are
made on the phasmid fauna from other geographical regions, numerous further unpublished
synonyms will be identified. A detailed background on the Brunner von Wattenwyl collection
in Vienna is included in Brock (in press).
References
Brock, P.D. (in press) Catalogue of type specimens of stick and leaf-insects in the Naturhistorisches Museum Wien
(Insecta: Phasmida). Annalen des Naturhistorischen Museums in Wien, Serie B.
Brunner von Wattenwyl, K. (1907) Die Insektenfamilie der Phasmiden. Volume 2. Verlag Engelmann, Leipzig.
Carl, J. (1913) Phasmides nouveaux ou peu connus du Museum de Geneve. Revue Suisse de Zoologie, 21(1): 1-57.
Colenso, W. (1885) A Description of some newly discovered New Zealand Insects believed to be new to Science.
Transactions of the New Zealand Institute, Zoology, 17: 151-155.
Hutton, F.W. (1898) The Phasmidae of New Zealand. Transactions of the New Zealand Institute, Zoology, 30: 160-
166.
Hutton, F.W. (1899) Revision of New Zealand Phasmidae. Transactions of the New Zealand Institute, Zoology, 31:
50-59.
Phasmid Studies, 6(1): 22
Taxonomic changes to New Zealand stick insects
Ragge, D.R. (1965) Grasshoppers, Crickets and Cockroaches of the British Isles. F. Warne & Co., London.
Redtenbacher, J. (1906) Die Insektenfamilie der Phasmiden. Volume 1. Verlag Engelmann, Leipzig.
Redtenbacher, J. (1908) Die Insektenfamilie der Phasmiden. Volume 3. Verlag Engelmann, Leipzig.
Salmon, J.T. (1954) A New Genus and Species of Phasmidae from New Zealand. Transactions of the Royal Society
of New Zealand, 82(1): 161-168, plates 7-9.
Salmon, J.T. (1991) The Stick Insects of New Zealand. Reed Books, Auckland.
Stal, C. (1875) Recensio Orthopterorum, Revue critique des Orthopteres decrits par Linne, de Geer et Thunberg.
Volume 3: 4-105. P.A. Norstedt & Sdner, Stockholm.
Westwood, J.O. (1859) Catalogue of Orthopterous Insects in the collection of the British Museum. Part I.
Phasmidae. British Museum, London.
White, A. (1846) The Zoology of the Voyage ofH.M.S. Erebus and Terror, 1. Insects of New Zealand. E.W.Janson,
London pp. 1-27.
PRAYING MANTIDS
The Mantis Study Group was
formed in 1996 to encourage the
study and rearing of praying
mantids. It produces a quarterly
newsletter containing a variety of
articles.
For further information contact:
The Membership Secretary,
Paul Taylor,
24 Forge Road,
Shustoke,
Coleshill,
Birmingham,
B46 2AU,
U.K.
Phasmid Studies, 6(1): 23
A glossary of terms used to describe phasmids
P.E. Bragg, 51 Longfield Lane, Qkeston, Derbyshire, DE7 4DX, U.K.
Abstract
A brief glossary of descriptive terms used for external morphology of phasmids, including synonyms. Particular
attention is paid to terms which are specific to phasmids or are used in a restricted sense when applied to phasmids.
Some features are illustrated.
Key words
Phasmida, Glossary, Terminology, Morphology.
Introduction
I have received several requests for an article explaining the terminology that is used in
descriptions of phasmids. For those without access to a good entomological dictionary, or
a morphological textbook, the terminology can be confusing. In the past, authors have used
terminology from various sources when describing phasmids, and although many terms have
a meaning which applies equally to any insect, some terms have a specific or restricted
meaning when applied to phasmids. Generally authors have used terminology which is
usually applied to Orthoptera, but the structures may differ from those in Phasmida. Some
terms are used differently by different authors (e.g. use of the word wing is often restricted
to the hindwing, with elytron being used for the fore wing, but some people use wing for both
hindwing and forewing. To avoid confusion in this particular case it is best if elytron and
hindwing are used, and the term wing avoided whenever possible). Some terms are
hyphenated by some authors and not by others, it is best to avoid using unnecessary hyphens:
i.e. use hindwing or hind wing and mid leg or midleg, but not hind-wing or mid-leg.
An English language glossary of phasmid terminology has never been published, and
some of the terms used in the past have been inadequately defmed, or usage has changed.
It should be recognised that this article is intended only as a brief guide for those without
easy access to more thorough texts. Two of the most widely available texts on morphology
are Snodgrass (1935) and Chapman (1982); a very widely available book containing a section
on morphology is A general textbook of entomology by Imms (various editions, 1925-1994).
An illustrated glossary was recently published in French (Lelong, 1996a, 1996b, 1996c).
A number of entomological dictionaries have been published but they are generally not
widely available. I have three entomological dictionaries and offer the following comments
on them:
The Dictionary of Entomology by N.K. Jardine (1913), Janson & Sons, London; 259 pages.
Veiy useful, particularly when working with old descriptions, but obviously lacking
modem terms. Occasionally available second hand.
A Dictionary of Entomology by A. W. Leftwich (1976), Constable & Company, London; 360
pages; ISBN 0-09-460070-8. This is not a dictionary: it is an entomological
encyclopedia which contains many family, generic and common names of insects, and
very few descriptive terms. I do not recall ever having used it as a dictionary.
Occasionally available second hand.
The Torre-Bueno Glossary of Entomology by S.W. Nichols (1989) New York Entomological
Society; 840 pages; ISBN 0-913424-13-7. This is a completely revised edition of A
Glossary of Entomology by J.R. de la Torre-Bueno (1937); 50 people are credited as
editorial contributors. It is invaluable, not only does it clearly define about 16000
terms but it also refers to alternatives and gives the source of the term. This book
should still be available from the publishers.
Phasmid Studies, 6(1): 24
Glossary of morphological terms
The following glossary is intended to include only those terms likely to be used in species
descriptions, it excludes most terms which are likely to be found in a general dictionary.
Some basic features not in the glossary are illustrated in figure 1 . Most of the terms apply
to adult insects but a few apply to eggs, for further egg terminology see Sellick (1992). In
many cases I have included some explanation with a basic definition. Where alternatives exist
they are indicated in square brackets following the definition; I have indicated reasons for my
personal preferences. Many terms are of Latin origin: some of these are commonly made
plural by the addition of the English endings s ot es, others are used with the Latin plural;
where the Latin plural is usually used this is given in brackets.
Terms used for the carinae of the legs, body surfaces, genitalia and the wing venation
are discussed in more detail following the alphabetical list.
Figure 1. Basic features of a typical phasmid.
Phasmid Studies, 6 ( 1 ): 25
Abdominal segments; The segments making up the abdomen are numbered from front to
back. Traditionally Roman numerals have been used but the use of Arabic numerals
is becoming common. The first, tenth and eleventh segments have specific names:
Median segment, Anal segment, and Lamina supraanalis.
Acanthotaxy: The naming of spines on the body and head. This is particularly useful for
members of the Heteropteryginae; spines have been described and illustrated by Rehn
& Rehn (1938).
Ala (Alae): The hindwing. It is attached to the anterior of the metanotum and is divided into
the leathery costal region and the folding anal region, [see Wing]
Anal process: A process projecting from, and part of, the anal segment; the processes are
usually paired although the two may be of unequal size (e.g. in Presbistus spp.).
Anal region: The folding part of a phasmid’s hindwing. This is often transparent or
translucent, and only rarely opaque.
Anal segment; The tenth abdominal segment. In most species this is the last full-sized
segment, but in females of some species the 11th segment may be large.
Appendicular ovipositor: An ovipositor formed by elongated ovipositor valves, it projects
beyond the end of the abdomen.
Archedictyon: A network of non-directional veins in the costal region of the wing or in the
elytron. It is these veins which make the elytron and costal region of the hindwing
thicker and stiffer than the anal region of the hindwing.
Areola; A sunken area on the ventral surface of the apex of the tibiae. This depression is
roughly triangular. This feature is used to split phasmids into two groups: the
Areolatae which have the sunken area, and the Anareolatae which do not.
Arolium (Arolia): A pad between the claws (ungues) of the pretarsus.
Binnenkorper: A hard sclerotization found in the genitalia of some female Asciphasmatinae;
may protrude from the left side of the operculum; the function is unknown.
Body length: The combined length of head, thorax and abdomen, including the operculum.
Some authors seem to have excluded the operculum from the body length, so where the
operculum extends beyond the end of the anal segment it is preferable to state clearly
whether the operculum is included in the body length.
Capitulum: A raised structure on the operculum of the egg (not present in all species).
Carina (Carinae): A ridge or raised line, typically along the leg, or centre line of the body.
Cercus (Cerci): Paired appendages attached to the anal segment. They may project beyond
the end of the abdomen or be hidden underneath; in a few cases they are enlarged leaf-
like structures.
Collar: The part of the capsule of an egg which surrounds the operculum; this is often a
narrowing, and occasionally an elongation, of the capsule.
Costal region: The anterior portion of the hindwing. This is thickened, opaque, and does
not fold; it is the part of the hindwing which is uppermost when the wings are folded.
Dentate: See toothed.
Dorso-anterior carina (carinae): A carina on the front, upper comer of the femur or tibia.
Dorso-posterior carina (carinae): A carina on the back, upper comer of the femur or tibia.
Elytron (Elytra): The fore wing. Attached to the posterior of the mesonotum. [Tegmen].
Foramen: see Pronotal foramen.
Gonapophysis (Gonapophyses): Appendages on the female’s abdomen which are usually
hidden by the operculum, but in some species they form part of an appendicular
ovipositor and are clearly visible (see fig. 5). [Ovipositor valves].
Granular: See Granulose.
Granulose: Covered in small granules; generally circular, and height is less than the
diameter. [Granular].
Phasmid Studies, 6 ( 1 ); 26
A glossary of terms used to describe phasmids
Hindwing: Attached to the anterior of the metanotum. The term hindwing (or hind wing,
or hind-wing) is usually used, the alternative, ala, is rarely used. [See Wing, Costal
region. Anal region].
Lamella (Lamellae): A thin sheet. Often used to refer to a thin sheet or lobe along the
Carina of a leg, typically running the whole length of the femur or tibia (frequently
found in Lonchodinae).
Lamina subgenitalis: The plate covering the genital opening. Rather an out-dated term.
[Subgenital plate; Operculum of female; Poculum of male] .
Lamina supraan^is: The 11th tergite (11th dorsal abdominal segment); absent in males but
can be seen in some females, it may be quite long but is more often minute or absent.
[Supra-anal plate].
Median segment: The first abdominal segment. This is fused with the metanotum to varying
degrees, in some instances it is indistinguishable.
Median sternite: The stemite of the median segment (indistinguishably fused with the
metastemum in most cases).
Median transverse groove: A indentation running across the pronotum, usually more or less
in the middle; it is particularly noticeable in some groups, e.g. Dares, but in many
groups can only seen with magnification. It marks the position of the sulcus, the joint
between the two plates which make up the pronotum. Sometimes just referred to as the
sulcus.
Medio- ventral carina (carinae): A carina in the middle of the ventral face of the femur or
tibia.
Mesonotum: The upper surface of the mesothorax.
Mesothorax: The second section of the thorax. The middle legs and elytra are attached to
the posterior of this segment.
Metanotum: The upper surface of the metathorax.
Metathorax: The third section of the thorax. The hindwings are attached to the anterior, and
the hind legs to the posterior of this segment.
Notum: A tergum of a thoracic segment, i.e. a dorsal plate of the thorax (see pronotum,
mesonotum, metanotum).
Opercular angle: The angle the operculum of an egg makes with the longitudinal axis. This
may be either negative or positive.
Operculum (Opercula): Used in two distinct contexts:
a) The lid of the egg.
b) Covering of genital opening of the females (the term has also used for males by
some authors but should be restricted to females; poculum should be used for males).
[Lamina subgenitalis; Subgenital plate].
Ovipositor: A device for egg laying, found in many species which push eggs into soil, cracks
or plant tissues. It is formed by either an elongated lamina supraanalis and elongated
operculum (an oviscapt); or by elongated valves and the operculum (an appendicular
ovipositor).
Oviscapt: An ovipositor which is formed by elongation of the operculum (8th stemite) and
lamina supraanalis (11th tergite); examples occur in Heteropterygini, Obrimini, some
Eurycanthinae, and some Necrosciinae. It is analogous to that found in several
Dipteran groups where the oviscapt is formed by a modified 7th stemite and 7th tergite.
Pectinate: Comb-like; the term is used to refer to the serrated ungues of Aschiphasmatinae.
Pleurite(s): A pleural sclerite, i.e. a lateral plate: one on the side of the body, between the
tergum and sternum. They are most commonly seen on the thorax and may be very
large and distinct in some groups, e.g. on the diorax of Heteropteryginae.
Phasmid Studies, 6 ( 1 ): 27
P.E. Bragg
Poculum (Pocula): The covering of the male genital opening; the 9th sternum. Often
referred to as the male operculum, but the use of poculum is preferable since it cannot
be confused with the female operculum. [Lamina subgenitalis; Subgenital plate].
Polar body: A mound on the egg at the opposite end to the operculum.
Praeopercular organ: An organ on the 7th stemite of females, used during copulation. It
is usually composed of one or more bumps, ridges or flaps. It is not present in all
species. [Preopercular organ].
Preopercular organ: See praeopercular organ.
Pretarsus: The distal segment of the tarsus, i.e. the claws (ungues) and arolium at the end
of the 5th tarsomere (3rd tarsomere in Timema),
Pronotal foramen: A distinctive opening on the anterior of the pronotum, used as a
diagnostic character of the tribes of Heteropteryginae (Rehn & Rehn, 1938).
Pronotum: The upper surface of the prothorax.
Prothorax: The first segment of the thorax. The fore legs are attached to this segment.
Punctate: Covered with many small pits (used in description of the surface structure of
eggs).
Rugose: Wrinkled; covered in small ridges.
Rugulose: Minutely rugose; fmely wrinkled.
Scabrous (or Scabrose): Rough; irregularly and roughly rugose; possessing short sharp
projections or wrinkles.
Sclerite: A hard plate of the body. These may be given more precise names depending on
where they are on the body, i.e. tergite, stemite.
Setose: Covered in setae; covered in stiff hairs.
Stemite: A ventral plate; a hardened plate which makes up part (or all) of a sternum.
Sternopleurite: A lateral plate on the ventral surface of the body, these are rarely evident
externally.
Sternum (Sterna): The ventral part of a segment, including stemites and stemopleurites.
Often stemopleurites are not evident, in which case the sternum may consist of a single
stemite.
Subgenital plate: A plate covering the genital area. [Lamina subgenitalis; Operculum,
Poculum] .
Supra-anal plate: The 11th abdominal segment. [Lamina supraanalis].
Tarsomere: A segment of the tarsus. There are five segments in all phasmids except
Timema which have only three.
Tegmen (Tegmina): The fore wing. Use of this term appears to have decreased, with a
corresponding increase in the term elytron in recent years.
Tergite: A dorsal sclerite; a hardened plate on the dorsal surface of the body which makes
up part (or all) of a tergum.
Tergum (Terga): The dorsal part of any segment. Although the term applies to all
segments, notum is often used to refer to the thoracic terga.
Tooth: A tubercule in which the height is greater than the diameter; a short blunt spine.
Toothed: Bearing numerous teeth. [Dentate] .
Tuberculate: Covered in tubercules. Finely tuberculate is generally considered the next
stage after granulose i.e. more strongly projecting than granulose.
Tubercule: A blunt or irregularly topped stmcture, not pointed like a spine; usually small but
the term is also used to refer to large wart-like stmctures; often of an irregular shape.
(See also Tooth and Vermcose).
Ungues: The claws of the pretarsus.
Phasmid Studies, 6 ( 1 ): 28
A glossary of terms used to describe phasmids
Ventro-anterior carina (carinae): A carina on the front, underside comer of the femur or
tibia.
Ventro-posterior carina (carinae): A carina on the back, underside comer of the femur or
tibia.
Verrucose: Covered in irregularly shaped lobes or wart-like protuberances.
Vomer: A moveable sclerotization on ±e 10th abdominal sternum of the male, usually more
or less triangular in shape; used during copulation. [Vomer subanalis].
Vomer subanalis: See Vomer.
Wing: The terms elytron or tegmen are almost always used when referring to the forewing
of a phasmid. The term hindwing is usually used in preference to ala. If the term mng
is used on its own it usually refers to the hindwing since in most phasmids the forewing
is relatively insignificant. [See Elytron, Tegmen, Ala, Hindwing]
The carinae of the legs
The surfaces and carinae of the legs are named on the following basis: with the leg extended
at right angles to the body, the upper surface is dorsal, lower surface is ventral, forward
facing surface is anterior, backward facing surface is posterior. Thus the four main carinae
are named: dorso-anterior, ventro-anterior, dorso-posterior, and ventro-posterior. A carina
between any of these carinae is termed a median carina and named according to the face on
which it is located, e.g. medio-ventral.
Some authors have used outer for dorsal, and inner for ventral: based on the flexed leg.
Since inner and outer could also be confused with inner meaning towards the body and outer
away from the body, it is best to avoid these terms.
Descriptions of surfaces
Surfaces are described using the following terms (in order of ascending roughness): Smooth,
Rugulose, Rugose, Scabrous, Verrucose; these terms are used when the whole surface is more
or less uniform. The terms Granulose and Tuberculate are also used to describe surfaces,
with granulose indicating an even covering of rounded granules, and tuberculate indicating
a surface with some rough projections which may range from sharp granules to isolated
verrucose structures; there is an overlap between tuberculate and scabrous and verrucose but
the latter two terms are used only if the whole surface is covered in irregularities, a
tuberculate surface clearly has flat areas between the individual tubercules.
Individual projections are referred to using the terms Granule, Tubercule, Tooth, Spine.
Sizes of spines present a problem because what one person considers small another may
consider large. Generally spine sizes are given relative to the other spines on the body. The
general nature of the genus, tribe or family group under consideration will influence the terms
used: a spine termed minute in the very spinose Heteropteryginae may be rated medium in
a species of a relatively spineless group such as Aschiphasmatinae. To try to be consistent,
I use the following as a rough guide for my own descriptions: minute (microscopic); small
(just visible to the naked eye); medium (clearly visible); large (very obvious); very large, and
extremely large (usually at least as high as the body).
Wing venation
Ragge (1955) surveyed the wing venation of the Phasmida, and applied the Comstock-
Needham terminology to the veins. Hamilton, in his study of insect wing venation
(Hamilton, 1971; 1972a; 1972b; 1972c) states that the radial and sector are fused and that
Ragge had been unaware of this and had therefore used the wrong terms for a number of the
veins. Table 1 gives the equivalent terms used by Ragge and Hamilton.
Phasmid Studies, 6 ( 1 ): 29
P.E. Bragg
The areas of the wing are named after the vein immediately anterior to the area, i.e.
the area behind the subcostal vein is the subcostal area, that behind the radial vein is the
radial area, etc. The main use of wing venation in phasmid descriptions is to name areas of
the wing for the purpose of describing the wing coloration.
The elytron of phasmids is usually reduced to a rudimentary flap, or is absent. In
species which have reduced wings, the wings and elytra may be of similar size and the
elytron may be much wider than the costal region of the wing.
The hind wings are divided into two distinct regions, the costal region and the anal
region. The costal region extends from the costal vein to about the empusal vein, and is
thickened and usually coloured. The radial vein is usually very obvious as the main vein of
the costal region. The anal region is thin, folding, and usually either colourless or
translucent, but in some species the anal region is strongly coloured. The anal veins arise
from two distinct points and form two sets, and A 7 onwards; the exact number of anal
veins in a species is difficult to assess without fully spreading the wing which usually
necessitates removing it.
The Phylliidae are atypical phasmids: in females the elytra are large while the wings
are absent; in males the wings and elytra are similar to typical phasmids except that the costal
region of the wing is not coloured.
Ragge (1955)
Hamilton (1971-72)
Costa
C
Costa
C
Subcosta
Sc
Subcosta
Sc
Radial
R
Radial
R
Sector (fused to R)
S
Radial sector
Rs
Media
M
Media (anterior)
Cubitus
Cu,
Media (posterior)
Mp
Cubitus
Cuj
Cubitus
Cu
Plical
P
First anal
Ai
Empusal
E
Anal
A2-7
Anal
A1.6
Table 1 Equivalent names used for veins by Ragge and Hamilton.
External genitalia of phasmids
Two features of the genitalia: the praeopercular organ of females, and the vomer of males,
are unique to Phasmida, although they are not found throughout the order. The male
genitalia was discussed by Snodgrass in his review of the genitalia of Orthopteroid insects
(Snodgrass, 1937), but phasmids were not mentioned by Scudder in his paper on the
comparative morphology of the insect ovipositor (Scudder, 1961). Gunther (1970) in Tuxon’s
Taxonomist's glossary of genitalia in insects makes only a fairly brief mention of the male
and female genital anatomy and does not distinguish between the three distinct types of
ovipositor.
The anal segment bears a pair of cerci which are typically cylindrical or slightly
Phasmid Studies, 6(1): 30
A glossary of terms used to describe phasmids
conical. The cerci may be straight or incurving and the apex may be smoothly rounded or
may bear a small spine. In some species the cerci have a complex shape; leaf-like cerci
occur in some groups.
Figures 2-5.
2. Poculum.
3. Scoop-shaped ovipositor with deep keel.
4. Oviscapt.
5. Appendicular ovipositor.
Phasmid Studies, 6 { 1 ): 31
Male genitalia
The male genitalia are covered ventrally, and usually also laterally, by the 9th sternum (fig.
2) which has been variously termed the lamina subgenitalis, operculum, and poculum; the 9th
sternum is usually divided into two stemites, the posterior of which has been termed the
subgenital lobe (Snodgrass, 1937), The term poculum was apparently first used by Key
(1970) for just the posterior stemite, but in Nichols (1989), for which Key was an editorial
contributor, it refers to the whole of the 9th sternum and I have followed that interpretation
here (and elsewhere). Poculum should be used in preference to the alternatives in order to
avoid possible confusion with the operculum (8th sternum) of the female.
In some species there is a large sclerotized organ, the vomer, which is pushed into the
praeopercular organ of the female. In some species, particularly in Lonchodinae the anal
segment is deeply cleft, forming a pair of claspers which are used to grip the female.
Female genitalia
The female genitalia are covered ventrally and laterally by the 8th sternum which has been
termed the lamina subgenitalis or sub genital plate, or operculum; the term operculum is now
almost invariably used. In some groups e.g. Heteropteryginae, Phasmatidae and
Lonchodinae, there is often a structure on the posterior of the 7th sternum which is termed
the praeopercular organ and varies from a small hollow, a rounded bump, a spine, to a pair
of large leaf-like structures. The praeopercular organ is gripped by the male during
copulation and appears to serve as a guide to positioning of the male’s genitalia. The internal
organs which may be visible externally comprise three pairs of valvulae, or gonapophyses.
In some species of the genus Presbistus Kirby (Aschiphasmatinae) there is a shiny black
sclerotized organ, termed the binnenkdrper by Gunther (1933; 1970), which may protrude
from the right side of the genital opening; the function of this organ is unknown.
There are three distinct forms of ovipositor in Phasmida. The typical form is a scoop-
shaped operculum varying in depth from an almost flat plate in some Necrosciinae and
Aschiphasmatinae to a deep scoop such as that found in most Lonchodinae; the deep forms
often have a distinct ventral or apical keel (fig. 3). The operculum usually reaches almost
to the end of the abdomen but in some genera or species it projects well beyond the end.
The second form of ovipositor is the oviscapt, formed by elongation of the operculum
and the 11th tergite (fig. 4); it is analogous with the structure found in several Dipteran
groups formed by a modified 7th stemite and 7th tergite. An oviscapt occurs in some
Heteropteryginae, some Eurycanthinae, some Necrosciinae, and in at least one "undescribed"
South American genus of Heteronemiinae (Gunther, 1970: 60).
The third form, termed an appendicular ovipositor, is formed by an elongated and
laterally compressed operculum and elongated gonapophyses; it is not homologous with the
typical orthopteran appendicular ovipositor which is formed from the gonapophyses alone.
An appendicular ovipositor is distinguished by the gonapophyses clearly projecting beyond
the anal segment (fig. 5). To be strictly correct the structure found in these phasmids should
be termed a semi-appendicular ovipositor because the operculum is not an appendage, but
since there are no known phasmids with a tmly appendicular form the more convenient term
appendicular can be safely applied. An appendicular ovipositor is found in a group of closely
related genera of Necrosciinae: Centrophasma, Diardia, Diesbachia, Galactea, Orxines,
Parastheneboea.
References
Chapman, R.F. (1982) The insects: structure and function. [3rd ed.] Hodder & Staughton, London.
Gunther, K. (19?3) Uber eine kleine Sammlung von Phasmoiden und Forficuliden aus Melanesien. Verhandlungen
der Natujforschenden Gesellschaft in Basel 44(2): 151-164.
Phasmid Studies, 6(1): 32
A glossary of terms used to describe phasmids
Gunther, K. (1970) Chapter 12, Cheleutoptera (Phasmoidea) pp. 58-61, in Tuxon, S.L. Taxonomist’s glossary of
genitalia in insects. [2nd edition] Munksgaard, Copenhagen.
Hamilton, K.G.A. (1971) The Insect Wing, Part I. Origin and development of wings from notal lobes. Journal of
the Kansas Entomological Society, 44(4): 421-433.
Hamilton, K.G.A. (1972a) The Insect Wing, Part II. Vein homology and the archetypal insect wing. Journal of the
Kansas Entomological Society, 45(1): 54-58.
Hamilton, K.G.A. (1972b) The Insect Wing, Part III. Venation of the orders. Journal of the Kansas Entomological
Society, 45(2): 145-162.
Hamilton, K.G.A. (1972c) The Insect Wing, Part IV. Venational trends and the phylogeny of the winged orders.
Journal of the Kansas Entomological Society, 45(3): 295-308.
Imms, A.D. (1925-1994) A general textbook of entomology . London. [Many revised editions].
Key, K.H.L. (1970) Phasmatodea (Stick-insects) in CSIRO, The Insects of Australia, volume 1. pp. 348-359.
Lelong, P. (1996a) Le dictionnaire des phasmes. Le Monde des Phasmes, 32: 15-20.
Lelong, P. (1996b) Le dictionnaire des phasmes (suite). Le Monde des Phasmes, 33: 16-26.
Lelong, P. (1996c) Le dictionnaire des phasmes (suite). Le Monde des Phasmes, 34: 17-25.
Nichols, S.W. (1989) The Torre-Bueno Glossary of Entomology Revised edition). New York Entomological Society.
Ragge, D.R. (1955) The wing-venation of the order Phasmida. Transactions of the Royal Entomological Society of
London, 106: 375-392.
Rehn J.A.G. & Rehn J.W.H. (1938) The Orthoptera of the Philippine Islands, Part 1. - Phasmatidae; Obriminae.
Proceedings of the Academy of Natural Sciences of Philadelphia, 90: 389-487.
Scudder, G.G.E. (1961) The comparative morphology of the insect ovipositor. Transactions of the Royal
Entomological Society of London, 113(2): 25-40.
Sellick, J. (1992) The phasmid egg. Phasmid Studies, 1(1): 8-9.
Snodgrass, R.E. (1935) Principles of Insect Morphology . McGraw-Hill, New York.
Snodgrass, R.E. (1937) Male genitalia of Orthopteroid insects. Smithsonian Miscellaneous Collection, 96(5): 107
pages [Phasmatodea, pp. 23-29].
Phasmid Studies, 6(1): 33
Reviews and Abstracts
Book Review
A guide to the stick and leaf insects of Singapore by Francis Seow-Choen (1997), with
foreword by David Rentz. Published by The Singapore Science Centre, Singapore. Softback,
160 pages, 122 colour illustrations, 5 black and white figures (10cm x 15cm), Price S$5. 15.
ISBN 981-00-8628-8. Reviewed by Paul D. Brock.
This is an excellent little colour pocket book with an attractive leaf-insect design on the
cover, part of a popular, inexpensive series of nature books published by the Singapore
Science Centre. Several years ago little was known about phasmids from Singapore, apart
from mainly historic records. The author has succeeded in making a thorough survey of the
Singapore phasmid fauna, collecting widely and listing previously unrecorded foodplants; no-
one has done more to popularise the study of phasmids in Asia.
The book amply succeeds in its aim of educating the public of Singapore, by using some
general introductory sections on these fascinating insects e.g. anatomy, functions, defence and
reproduction. The main section gives a brief description and notes on each species found in
Singapore, with many colour photographs of the adults and eggs (where known), or
occasionally sketches. Common names have been used, which should prove popular with
non-specialists. All but one of the 40 species included are illustrated (including three species
being described in my forthcoming book on the phasmids of Peninsular Malaysia and
Singapore).
This book is well printed, with good quality photographic reproductions, which often
show insects on their foodplants in nature, at night; rarely featured in previous publications
on phasmids. A very short glossary of some terms is included, in addition to notes on
phasmid groups and selected references.
To sum up, this excellent, accurate pocket book will be a must for every phasmid
enthusiast and should considerably raise interest in phasmids in S.E. Asia, especially as many
of the species are also found in Peninsular Malaysia. Individuals living in a warm climate,
may even be tempted to build a ’hut’ in their garden to keep live insects (as illustrated),
alternatively they might venture out at night with torchlight to look for these insects! With
so much crammed into this book, the fact that information on the wider distribution of species
is not always mentioned is not unreasonable, however, the mention of a number of non-
Singaporean species, sometimes without any reference to their distribution, may be confusing
to the non specialist. For the benefit of the specialist, there are a few name variations,
ignoring those currently in press: Lonchodes geniculatus Gray, should not have the author’s
name in brackets; in the case of Bacteria ridleyi, the generic name should read Bactricia;
Gargantuoidea pkaetusa (Westwood) is the valid name, rather than G. gargantua; Datames
mouhotii should be spelt as shown, with an extra ’/’; the male of Planispectrum bengalensis
was described in 1995, and Presbistus flavicomis is associated with Singapore from a historic
record, but not mentioned.
PSG members interested in obtaining this book, please send s.a.e. for details of price
and ordering arrangements to P.D. Brock, "Papillon", 40 Thorndike Road, Slough, SL2 ISR,
or await details in the September PSG Newsletter. A bulk order will reduce costs
considerably.
Phasmid Studies, 6(1): 34
Phasmid Abstracts
The following abstracts briefly summarise articles which have recently appeared in other
publications; also included are publications published since 1992 (the first issue of Phasmid
studies) which have only recently come to the attention of the editor. Some of these may be
available from local libraries. Others will be available in university or college libraries,
many of these libraries allow non-members to use their facilities for reference purposes free
of charge. In the UK libraries can usually obtain publications which they do not hold by
using the inter-library loan system; there is usually a charge for this service. A similar inter-
library loan system operates in many other countries.
The editor of Phasmid Studies would welcome recent abstracts from authors so that they may
be included in forthcoming issues. In the case of publications specialising in phasmids,
Phasma and Le Monde des Phasmes, only the longer papers are summarised.
Ali, D.W. & Orchard, I. (1996) Inununohistochemical localization of tyrosine hydroxylase
in the ventral nerve cord of the stick insect, Carausius morosus, including neurons innervating
the salivary glands. Cell & Tissue Research, 285(3): 453-462.
The distribution of tyrosine hydroxylase-like immunoreactive neurons is mapped in the
ventral nerve cord of the stick insect, Carausius morosus. This study also examines the
tyrosine hydroxylase- and serotonin-like immunoreactive elements in the salivary glands of
Carausius morosus. Tyrosine hydroxylase is the first and rate-limiting enzyme in the pathway
for the production of catecholamines; therefore, tyrosine hydroxylase-like immunoreactive
neurons are likely to contain catecholamines. Approximately 225 tyrosine hydroxylase-like
immunoreactive neurons are present in the ventral nerve cord. The majority of these neurons
appear to be intemeurons. The suboesophageal ganglion contains the only unpaired neuron
and the only pair of peripherally projecting tyrosine hydroxylase-like immunoreactive neurons
in the ventral nerve cord. The peripherally projecting neurons project to the salivary glands
via the salivary nerve. Each neuron in this pair is termed the salivary neuron 1. The
remaining tyrosine hydroxylase-like immunoreactive neurons in the ventral nerve cord are
intemeurons and exhibit a characteristic distribution within the thoracic and the abdominal
ganglia. Serotonin-like immunoreactivity is also present in the salivary glands. Positive
staining of the suboesophageal ganglion for serotonin-like immunoreactivity indicates the
presence of several neuron pairs including a large pair along the ventral posterior midline that
project to the salivary glands via the salivary nerve. Each neuron in this pair is termed the
salivary neuron 2. Backfilling of the salivary nerve with cobalt chloride reveals the presence
of only two neurons within the suboesophageal ganglion that project to the salivary glands;
these neurons are the salivary neurons 1 and 2. Reverse-phase high-performance liquid
chromatography coupled with electrochemical detection of ventral nerve cord and salivary
gland homogenates confirms the presence of dopamine and serotonin.
Ansorge J. (1996) On the systematic position of Schesslitziella haupti Kuhn 1952 (Insecta:
Phasmatodea) from the Upper Liassic of northern Franconia (Germany). Palaeontologische
Zeitschrift, 70(3-4): 475-479.
The holotype of Schesslitziella haupti Kuhn, 1952 from the Upper Liassic of northern
Franconia is redescribed. The species is considered a representative of the Chresmodellinae
(Phasmatodea: Aerophasmatidae). Chresmodella Bode, 1953 is a younger synonym of
Schesslitziella Kuhn, 1952. [In German]
Phasmid Studies, 6 ( 1 ): 35
Phasmid abstracts
Baessler, D., Bueschges, A., Meditz, S. & Baessler, U. (1996) Correlation between muscle
structure and filter characteristics of the muscle-joint system in three orthopteran insect
species . Journal of Experimental Biology , 199( 10): 21 69-2 183.
In orthopteran insects, neural networks for joint control exhibit different characteristics
due to behavioural specializations. We investigated whether these differences are generated
purely by the neuronal networks, or whether characteristics of the muscles or joint
architecture (muscle-joint system) are also involved in these behavioural specializations. We
compared the properties of the muscle system moving the femur-tibia joint of the middle and
hind leg of three species, Carausius morosus, Cuniculina impigra and Locusta migratoria.
Four aspects were analyzed for the tibial extensor muscle: (i) the frequency-dependence of
motoneuronal activity in response to sinusoidal stimulation of the femoral chordotonal organ
(fCO), (ii) the muscle structure, (iii) the innervation pattern of the muscle and (iv) the
histochemical properties of the muscle fibres. These aspects were compared with the filter
characteristics of the open-loop femur-tibia control system and of the muscle-joint system
involved. Whereas in both phasmid species {Carausius morosus and Cuniculina impigra) the
motoneuronal activity steadily increases with sinusoidal stimulation of the fCO in the
frequency range 0.01-5Hz, in Locusta migratoria there is a decrease in motoneuronal activity
between 0.01 and 0.3Hz. The muscle structure is basically similar in all three species, as the
number of singly innervated muscle fibres (supplied by the fast extensor tibiae motor neurone,
FETi) decreases from proximal to distal. The number of triply innervated fibres supplied by
the FETi, the slow extensor tibiae (SETi) and the common inhibitor 1 (CI-1) is maximal in
the middle of the muscle, and the number of dually innervated fibres (supplied by SETi,
CI-1) increases from proximal to distal. Differences between the locust and the two phasmid
species exist in the distal portion of the muscle. The phasmid extensor tibiae muscle contains
a morphologically distinct bundle of muscle fibres, not present in the locust, which is mostly
dually innervated and which is larger in Cuniculina impigra. Similar results were obtained
for the histochemical characterization of the muscle fibres as revealed from their staining for
myofibrillar ATPase activity. The number of histochemically identified fast fibres decreased
from proximal to distal, while the number of slow fibres increased. In Carausius morosus
and Locusta migratoria, the percentage of slow fibres increased by up to 60-70% at the distal
end, while this increase was to almost 100% in Cuniculina impigra. Apparently, the larger
this distal region and the higher the percentage of slow, dually innervated fibres in it, the
lower is the upper comer frequency (the stimulus frequency at which the joint control system
produces a movement with 70% of its maximal response amplitude) of the muscle-joint
system. In summary, it appears that the upper comer frequency of the open-loop system in
Locusta migratoria (It 0.05Hz) results at least in part from properties of the neuronal joint
control network, but in Carausius morosus (0.5-1.0Hz) and Cuniculina impigra (0.1-0.2Hz)
it results from the upper comer frequency of the muscle-joint system.
Baessler, U. & Stein, W. (1996) Contributions of stmcture and innervation pattern of the
stick insect extensor tibiae muscle to the filter characteristics of the muscle-joint system.
Journal of Experimental Biology , 199(10): 2185-2198.
It is shown that the low-pass filter characteristics of the muscle-joint system of the
femur-tibia joint of the stick insect Cuniculina impigra result from co-contraction of the
extensor and flexor tibiae muscles. The most distal region of the extensor muscle, which
contains a high percentage of slow muscle fibres, is involved in this co-contraction. This
conclusion results from the following evidence, (1) Inertial and friction forces do not affect
the characteristics of the low-pass filter of the muscle-joint system. (2) There is some co-
contraction of the extensor and flexor muscles during sinusoidal stimulation of the femoral
Phasmid Studies, 6 ( 1 ): 36
Phasmid abstracts
chordotonal organ at high stimulus frequencies. Both muscles generate tonic forces that
increase with increasing stimulus frequency and also increase with time from the beginning
of stimulation until a plateau is reached. (3) For the extensor muscle, this tonic force is
produced by its most distal portion only. (4) Electrical stimulation of the common inhibitory
motoneurone (CI-1) reduces the tonic force generated in this most distal portion of the
extensor muscle. Therefore, CI-1 stimulation reduces the amplitude of tibial movement in
response to sinusoidal stimulation of the femoral chordotonal organ at stimulus frequencies
below 0.5Hz (over this frequency range, the tibial movement amplitude is a function of the
force amplitude produced by the whole extensor muscle and there is no co-contraction), but
at chordotonal organ stimulus frequencies of IHz and above, CI-1 stimulation increases the
tibial movement amplitude (in this case, movement amplitude is limited by the degree of
cocontraction of the extensor and flexor muscles). With repeated chordotonal organ
stimulation at higher stimulus frequencies, the tibial movement amplitude steadily decreases.
This must be a consequence of increasing levels of co-contraction of the extensor and flexor
muscles, since at low stimulus frequencies (no cocontraction) there is no reduction in
movement amplitude during repeated stimulations. It is concluded that co-contraction of the
extensor and flexor tibiae muscles prevents instability in the reflex loop in spite of the high
gain necessary for the generation of catalepsy. Therefore, the mechanism described can be
considered to be an adaptation to the ecological niche occupied by this animal. The
contribution of the distal part of the extensor muscle to this system can be switched off by
the CI-1 during active movements.
Baez, M. (1996) Nuevas citas de insectos en las Islas Canarias (Phasmatodea, Lepidoptera,
Embioptera). Boletin de la Asociacion Espanola de Entomologia, 20(1-2): 252-253.
In 1926 Bolivar included the Canary Islands in the distribution of Clonopsis gallica
(Charpentier) but no specific records were published. In 1982 Baez recorded a female
phasmid from La Metanza on the island of Tenerife but it was not certain that it was from
an established colony, the possibility of it being a casual introduction could not be discounted.
In 1994 the author collected two more female specimens from the same locality, confirming
this species does occur on Tenerife.
Bland, R.G., Gangwere, S.K. & Morales Martin, M. (1996) An annotated list of the
Orthoptera {sens, lot.) of the Canary Islands. Journal of Orthoptera Research, 5: 159-173.
The distribution of 117 Canary Island orthopteroid species belonging to Orders
Blattaria, Mantodea, Orthoptera, and Phasmida are presented based on the authors’
collections, museum specimens, and literature since the last list in 1954. The number of
species in each order and the percentage endemic to the archipelago are: Blattaria 24 (50%);
Mantodea 9 (67%); Orthoptera 83 (37%), and Phasmida 1 (0%). The same for families of
Orthoptera follows: Acrididae 41 (41%); Gryllidae 18 (17%); Gryllotalpidae 2 (0%);
Pamphagidae 4 (100%); Pyrgomoiphidae 1 (0%); Tetrigidae 1 (0%); and Tettigoniidae 16
(44%). Orthopteroid species diversity and the number of endemics were greatest on Tenerife
(82 species, 24 endemics), followed by Gran Canaria (64 species, 17 endemics), and La
Gomera (49 species, 14 endemics); Fuerteventura had the fewest number of species (28) and
the lowest number of endemics (5). Tenerife supported the highest number of single-island
endemics (8) and La Palma had the lowest (1). The highest percentages of endemics, 27%
to 29%, occurred on Tenerife, La Gomera, La Palma, Lanzarote, and Gran Canaria;
Fuerteventura had 18% and El Hierro 17%.
New species, changes in nomenclature, and unconfirmed records are discussed, as are
taxonomic problems encountered in taxa of the acridid genera Sphingonotus and Acrotylus.
Phasmid Studies, 6 ( 1 ): 37
Phasinid abstracts
Chen, S.C. & He, Y.H. (1996) A new species of Cnipsus from Yunnan, China (Phasmida:
Phasmatidae). Acta Entomologica Sinica, 39(3): 286-288.
This paper describes a new species of the genus Cnipsus from Yunnan Province. The
type specimen of Cnipsus colorantis n.sp. is deposited in the Institute of Zoology, Academia
Sinica. All the measurements in descriptions are in millimetres. [In Chinese]
Chen, S.C. & He Y.H. (1996) Description of two species of males of Parasipyloidea and
Trachythorax (Phasmatodea: Heteronemiidae) unknown before. Forest Research, 9(6):
664-665.
In this paper two new species Parasipyloidea emeiensis Chen & He and Trachythorax
fuscocarinatus Chen & He are described from males. The specimens are kept in the Insect
Collection of Beijing Forestry University. All measurements in descriptions are in mm. [In
Chinese]
Doerr, H., Hess, D. & GramoU, S. (1996) Interstitial voltage and potassium concentration
in the mesothoracic ganglion of a stick insect at rest and during neuronal activation.
Journal of Insect Physiology , 42(10): 967-974.
In the mesothoracic ganglion of the stick insect Cuniculina impigra the interstitial
voltage and potassium concentration were measured. The interstitial voltage was measured
using conventional glass microelectrodes and had a mean value of +17.6mV (± 4.7mV, N
= 14 animals). The potassium concentration in the interstitium was measured using ion-
selective microelectrodes and had a mean value of 4.3mM (+ l.lmM, N = 13 animals).
The interstitial voltage and the interstitial potassium concentration changed when the ganglion
was superfiised with solutions containing different potassium concentrations: in bath solutions
containing, e.g. high potassium concentrations, the interstitial voltage increased and the
interstitial potassium concentration increased. The interstitial voltage decreased by l-5mV
when neuronal activity was evoked by stimulating a nerve electrically. When an animal was
activated by stimulation with a paintbrush, the interstitial voltage decreased by 2.7mV (±
l.lmV, N = 3 animals) and the interstitial potassium concentration increased by 0.4mM (±
0.2mM, N = 3 animals).
Frantsevich, L.I. (1997) Artifactual motility of the subcoxal axis in a model insect leg with
skew joint axes. Journal of Theoretical Biology, 184(3): 111-111.
Kinematic reconstructions of joint angles during walking in stick insects revealed that
the subcoxal axis (SCA) was neither firm during the step (it was movable), nor was its
position matched to the line between articulation condyles. It has been demonstrated on
kinematic models of a leg that the application of a simplified leg model for reconstruction
caused an artifactual result: apparent movement of the SCA and its drift from the real firm
position. Simplified models assume that the plane where leg segments lie rotates about the
SCA or at least the axis of joint rotation is perpendicular to the leg segment. Real legs have
joints with non-parallel or non-orthogonal axes. Models considered in this article had an
oblique arrangement of joint axes. Real or artifactual motility of SCA is discussed.
Hennemaim, F.H. & Conle, O.V. (1996) Symetriophasma brevitarsa n.gen., n.sp. - eine
neue Phasmide aus Neuguinea (Phasmatodea: Eurycanthinae). Entomologische Zeitschrift,
106(11): 457-460.
Symetriophasma brevitarsa n.gen. n.sp., a new genus and species of Phasmatodea from
New Guinea, belonging to the subfamily Eurycanthinae, is described and figured. The
systematic position of the genus within the subfamily is briefly discussed. A second species.
Phasmid Studies, 6 ( 1 ): 38
Phasmid abstracts
Trapezaspis echinata Gunther, 1936, is also transferred to the new genus.
Hennemaiin, F.H., Conle, O.V. & Briickner, M. (1996) Dajaca nigrolineata n.sp. - eine
neue Phasmide aus Myanmar (= Burma), mit Bermerkungen zu dem Genus Dajaca Brunner
1893 (Phasmatodea: Phasmatidae: Aschiphasmatinae). Entomologische Zeitschrift, 106(8):
329-335.
A new species of Phasmida - Dajaca nigrolineata n.sp. - from Myanmar is described.
It is the first non-Bomean representative of the genus Dajaca Brunner, 1893. A review of
the genus is given, with a key to its species. The new species differs from D. monilicomis
Redtenbacher in the males being completely wingless. It is nearly related to D. jUiformis
Bragg. The eggs of D. monilicomis are described and illustrated for the first time.
Hennemann, F.H. & Conle, O.V. (1997) Die Gatmng Thaumatobactron Gunther, 1929 mit
der Beschreibung awei neuer Arten aus Papua Neuguinea (Phasmatodea, Eurycanthinae).
Mitteilungen aus dem Zoologischen Museum in Berlin, 73(1): 175-182.
Two new species of the genus Thaumatobactron Gunther, 1929 (T. granulosa n.sp. and
T. guentheri n.sp.) from Papua New Guinea are described and figured. A review of the
genus and a key to the males is given. Some information on the biology of the species is
included.
Hennemann, F.H. & Conle, O.V. (1997) Intraspezifische Variabilitat bei Lonchodes
femoratus (Stoll, 1787) nebst einigen Bemerkungen zu ihrer Synonymie (Phasmatodea:
Phasmatidae: Lonchodinae). Entomologische Zeitschrift, 107(1): 30-37.
In this paper the intraspecific variation of the phasmid Lonchodes femoratus (Stoll) is
discussed and illustrated. The material examined originates in a phasmid collection made by
P. Kibler in 1912 from the Key Islands, which is now housed in the collection of the
Naturkunde-Museum in Stuttgart (SMNS) and includes 158 females, 7 males and two large
female nymphs. The synonymy of the species is discussed and one new synonym {Prisomera
expulsum Brunner v. W., 1907) is listed, which is said to differ from L. femoratus by having
a struma on the mesonotum. The eggs are described and illustrated. [Editor’s note: The
correct name for L. femoratus is Lonchodes foliopeda (Olivier, 1792) - see Phasmid Studies,
4(1): 24 for details.]
Kittmann R., Schmitz J. & Bueschges A. (1996) Premotor intemeurons in generation of
adaptive leg reflexes and voluntary movements in stick insects. Journal of Neurobiology ,
31(4): 512-531.
We investigated the role of local nonspiking intemeurons involved in motor control of
legs in the stick insect, Carausius morosus. In a preparation that allowed the animals to
perform active leg movements such as adaptive tactile reflexes, proprioceptive reflexes, and
walking, we gathered the following results. Almost all tested nonspiking intemeurons that
provide synaptic drive onto motoneurons of the proximal leg muscles contribute to all of the
motor programs underlying tactile reflexes and voluntary leg movements such as walking,
searching, and rocking. Most of them are also involved in the generation of proprioceptive
reflexes. All motor programs for coactivation, avoidance reflexes, resistance reflexes, and
voluntary leg movements result from parallel pathways including nonspiking intemeurons that
support and others that oppose the motoneuronal activity. The contribution of a single
intemeuron to the different motor programs is specific: it can be supporting for one motor
program but opposing for the other. Even for the same motor program, for example,
coactivation, the contribution of an individual intemeuron can depend on the stimulus site
Phasmid Studies. 6 ( 1 ): 39
Phasmid abstracts
from where the response is elicited. Our results support the idea that the different motor
patterns for adaptive tactile reflexes, resistance reflexes, and voluntary leg movements emerge
from a multifunctional neuronal circuit that is reorganized corresponding to the motor
behaviour performed. The actual motor pattern is then shaped by distributed information
processing in parallel supporting and opposing pathways.
Mantovani, B., Tinti, F. Barilani, M. & Scali, V. (1996) Current reproductive isolation
between ancestors of natural hybrids in Bacillus stick insects (Insecta: Phasmatodea).
Heredity, 77(3): 261-268.
Interspecific hybrids raise a variety of developmental, reproductive, and evolutionary
issues. In Sicily, geographically and chronologically distinct hybridizations between the
highly differentiated Bacillus rossius and B. grandii have produced hybridogenetic strains and
clonal paithenogenetic species. In northern Sicily, all-female populations of facultatively
parthenogenetic B. rossius and bisexual B. grandii benazzii co-occur and we could test their
current hybridization through electrophoretic marker analyses; control crosses with allopatric
males were also carried out. Hybrid female progeny percentages ranged from 0-74 being
fewer in egg batches laid by parthenogenetic mothers than in those of amphimictic females;
no difference was noticed between sympatric and allopatric pairs. F2 hybrids of both sexes
proved sterile; although some eggs started cleaving, no hemiclonal or clonal progeny hatched,
only rare androgenetics being obtained. In currently produced hybrids a complete disruption
of gametogenesis occurs, so that genetic constraints between parental taxa appear stronger
now than in the past, most likely the result of ancestor evolution.
Seow-Choen, F. (1996) The leaf insects of peninsular Malaysia. Nature Malaysiana, 21(3):
68-73.
A magazine article which briefly reviews the leaf insects known from West Malaysia.
The article includes 13 colour photographs of Phy Ilium and one photograph of an egg.
Seow-Choen, F. (1996) A colourful stick insect from Sarawak. Nature Malaysiana, 21(4):
116-119.
A magazine article discussing the brightly coloured stick insect Dajaca monilicomis
Redtenbacher, 1906; with seven colour photographs. The native foodplant is Tristania sp.
but Eucalyptus robusta is eaten in captivity.
Seow-Choen, F. (1997) The joy of stick-insects. Singapore Scientist, 81: 10-14.
A magazine article with 18 colour photographs of South East Asian phasmids.
Zompro, O. (1997) Hermarchus leytensis n.sp., eine neue Phasmide von den Philippinen.
Entomologische Zeitschrift, 107(1): 38-40.
A new phasmid (Phasmatodea: Phasmatidae: Phasmatinae: Hermarchini
[= Phamaciini]) from the Philippine island of Leyte and its egg is described and figured.
Hermarchus leytensis n.sp. is the first member of the genus Hermarchus Stal, 1875 recorded
from the Philippine islands.
Phasmid Studies. 6 ( 1 ); 40
The egg of Baculofractum insignis (Brunner)
J.T.C. Scllick, "Montemassi", 31 Regent Street, Kettering, Northants, NN16 8QG, UK.
Abstract
The capitulum of the egg of Baculofractum insignis described, and the egg illustrated. The unusual features of the
egg are discussed.
Key words
Baculofractum insignis, egg structure.
The genus Baculofractum Zompro, 1995 was erected for what was previously Carausius
insignis. The significant reason for this was the discoveiy that the male of this species was
winged, whereas in Carausius, and indeed in the whole of the Lonchodinae in which it was
placed, males are apterous. As a consequence the new genus was placed in Necrosciinae.
Zompro (1995) described both the male and the egg of this species for the first time.
However the egg was illustrated only by a lateral black-and-white photograph. Both
description and photograph lack the capitulum of this egg. Zompro *s description (translated
from the German) is: — "Dimensions (average of five eggs): length 4.4mm, width 2.95mm,
height 3.75mm. Dark brown; round, laterally bevelled, surface with more or less round
depressions that are enclosed by very short-bristly raised areas. Micropylar plate darker,
raised above the egg surface, laterally widened at the level of the micropyle. Operculum flat,
with a conical projection. "
It can be seen from Fig. 1 that this is accurate, as far as it goes. The bristles are
indeed very short (around 0.01mm long) and difficult to detect. Zompro suspects that these
may be lost in older eggs. I have recently been sent a number of eggs of this species by Wim
Potvin, one of which retains its capitulum. Zompro (personal communication, 1997) saw
several hundred eggs of this species without finding a single one which retained its capitulum.
However the position of the capitulum stalk is clearly visible in those opercula which have
Figure 1. Egg of Baculofractum insignis.
A. dorsal; B. lateral; C. internal micropylar plate; D. operculum.
Surface patterning shown on lateral view only.
Phasmid Studies. 6(2): 41
7T.C. Sellick
lost the actual capitulum. Whatever the function of a stalked capitulum, this species shows
more than most the fragility of its attachment to the operculum. This capitulum is so far
unique in its minute size (0.2mm diameter) and very fme stalk (0.06mm thick). It is pale
yellowish-brown and button-shaped with a central depression, very much like a capitulum of
a genuine Carausius.
The internal micropylar plate follows roughly the outline of the external plate, though
narrower, and it is closed. This is the first egg with this type of capitulum that I have found
with a closed internal plate. This egg therefore shows a combination of characters from the
two subfamilies and is a further indication for the need to reexamine relationships within this
family. Stalked button capitula have only previously been found in the Lonchodinae (all of
which have open plates with median lines) whilst the closed internal plate most closely
resembles that of Phaenopharos Kirby, 1904 of the Necrosciinae, though the capitulum of this
genus is not stalked.
References
Zompro, O. (1995) Baculofractum n.gen. — ein neues Genus der Phasmida. Entomologische
Zeitung, 105(24): 488-491.
Phasmid Studies, 6(2): 41-42. Published January 1998.
Phasmid Studies, 6 ( 2 ): 42
A new species of Phobaeticus Brunner von Wattenwyl, from the
Philippines (Phasmatidae)
Paul D. Brock, "Papillon", 40 Thorndike Road, Slough, SL2 ISR, UK.
Abstract
Phobaeticus lumawigi n.sp. is described from a single female collected from Luzon, Philippines and deposited in the
Natural History Museum, London. This species is distinguished from others in the genus by its unusual leg serration.
Phamacia rigida Redtenbacher, 1908 is returned to the genus Phamacia St41, 1877 (from Phobaeticus Brunner von
Wattenwyl, 1907).
Key words
Phasmida, Phobaeticus lumawigi n.sp., Philippines.
Introduction
Mr Ismael Lumawig (Manila, Philippines) recently made an interesting collection of stick-
insects from various parts of the Philippines, which have been sent to me for identification;
they include several large stick-insects from Mountain Province, North Luzon, all except one
identified as Phamacia ponderosa Stal, 1877, type species of Phamacia Stal, 1877. This
single female is described as the first Phobaeticus Brunner von Wattenwyl, 1907, species to
be recorded from the Philippines; other Asian representatives of this genus include the longest
known insects. The taxonomy follows that of Brock (1996), which corrected various errors
in the literature relating to ’giant’ stick-insects, designated a type species for Phobaeticus and,
in particular, pointed out that Phobaeticus includes species with either winged or wingless
males; females are always wingless. The genus belongs to the family Phasmatidae, subfamily
Phasmatinae.
Phobaeticus Brunner von Wattenwyl, 1907
Phobaeticus Brunner von Wattenwyl, 1907: 194. Type species: Phobaeticus sobrinus
Brunner von Wattenwyl, 1907: 184, pi. 7.1a (c?), lb (9); designated by Brock, 1996: 30
(type locality - Sumatra, Si-Rambe).
Phobaeticus lumawigi n.sp.
Holotype 9; Philippines, Mountain Province, North Luzon, VIL1996, leg. I. Lumawig
(BMNH, London).
Female: Elongate, completely brown, largely smooth, with serrate legs, including a few
larger dentations on upper third of all femora; end of anal segment sharply triangularly
incised.
Head: Segment marginally longer than wide, with indistinct dark lines from eyes to
back of head. Eyes large, brown. Antennae same shade as body, basal segment long and
thin, 2.5 times length of next segment; tips broken off (believed to exceed length of fore
femora originally).
Thorax: Pronotum shorter and narrower than head, with a central indentation; viewed
laterally, front and hind part of segment are slightly raised. Mesonotum long and slender,
almost 6.5 times length of pronotum, and tapered gently towards hind part of segment.
Metanotum much shorter than mesonotum.
Abdomen: About same width as thorax; ninth segment only half the length of eighth.
Anal segment slightly longer than ninth; with a bold triangular incision, although the extreme
margins are rounded and covered with small hairs; a central, rounded extension (the lamina
supraanalis) protrudes beneath the incision. Minute hairs present on abdomen, which is
otherwise smooth. Cerci medium sized, slender, tapering sharply to an almost pointed tip.
Phasmid Studies, 6(2); 43
Paul D. Brock
Figures 1 & 2. Phobaeticus lumawigi n.sp.
1, Dorsal view. 2. End of abdomen, lateral view.
Phasmd Studies, 6 ( 2 ): 44
A new species of Phobaeticus from the Philippines
Operculum long, projecting beyond end of anal segment; rather tapered to a narrow tip.
Distal end of underside of seventh segment split into two central, stout pointed extensions.
Legs: Numerous small serrations on femora and tibiae, typical of the genus (but
sometimes larger in other species). Ventro-anterior carina of fore femur with a medium sized
triangular spine-like lobe four-tenths from base, beneath the regular serrations which are more
tooth-like than those on mid and hind femora. Mid and hind femora with large black sub-
basal triangular spine-like lobe on dorso-anterior carina, followed by a smaller lobe after a
gap of approximately 6mm; a pair of large spine-like lobes are present on the ventral carinae
beneath the largest spine (left hind leg has been regenerated and is slightly shorter than
normal). Mid tibiae also with a few larger lobes, both apically and basally, and an apical
broadened crest at base of dorsal surface; features also present to a lesser degree on hind
tibiae- Tarsi long.
Measurements (left fore leg is missing; measurements of hind leg are taken from the
right leg): Body length 202mm, head 9mm, antennae 62mm, pronotum 7.5mm, mesonotum
48.5mm, metanotum 14mm (25mm, including first abdominal or median segment). Femora:
fore 62mm, mid 46mm, hind 55mm. Tibiae: fore 83mm, mid 50mm, hind 65mm.
Operculum 30mm, cerci 3mm.
Etymology: Named after the collector Ismael Lumawig, in recognition of his
considerable recent efforts and enthusiasm in collecting stick-insects, despite the presence of
venomous snakes in some localities!
Discussion
This insect, the first Phobaeticus species recorded from the Philippines, may be distinguished
from other taxa by the unusual spine formation on its legs. In the absence of a male, it is
not clear with which "group" within the genus this species is closely linked i.e. those with
winged or wingless males. The egg is not known, but the adult morphology is sufficient to
readily distinguish Phamacia from Phobaeticus, I am aware of a current study placing
considerable importance on eggs in Phobaeticus and related taxa; however, whilst the study
of eggs may assist in distinguishing closely related taxa, variation of eggs within a genus can
sometimes be striking and until the eggs of a number of species are known, it does not appear
practical to draw meaningful conclusions.
Since publication of my 1996 paper (which includes listings of Phamacia and
Phobaeticus species), I have been able to examine type material in the Naturhistorisches
Museum Wien; as a result, I here reinstate Phamacia rigida Redtenbacher, 1908: 453
(holotype locality - Sumatra; Mt Battak) in the genus Phamacia (automatically transferred to
Phobaeticus in my 1996 paper). It will undoubtedly be necessary to make fiirther changes,
following Brunner von Wattenwyl and Redtenbacher’ s difficulty in distinguishing between the
genera. However, this will require critical examination of all relevant taxa, comparing
specimens alongside each other to establish the correct synonymy.
Acknowledgement
I would like to thank Mr Ismael Lumawig for sending the specimen described.
References
Brock, P.D. (1996) Changes of taxonomy in giant stick-insects. Phasmid Studies, 5(1): 25-31.
Brunner von Wattenwyl, K. (1907) Die InsektenfamiUe der Phasmiden, 2: 181-338. Verlag Engelmann, Leipzig.
Redtenbacher, J, (1908) Die Insektenfamilie der Phasmiden, 3: 339-576. Verlag Engelmann, Leipzig.
Stal, C. (1877) Orthoptera nova ex Insulis Philippinis descripsit. Ofversight af Kongliga Vetenskaps-Akademiens
Fdrhandlingar, 34(10): 33-58.
Phasmid Studies, 6(2): 43-45. Published January 1998.
Phasmid Studies, 6(2): 45
Reviews and Abstracts
Book Reviews
Your First Stick Insect by David Alderton (1997), with photographs by the author, Nick
Baker and Paul Brock. Published by TFH Kingdom Books, Waterlooville. Paperback, 33
pages, 28 colour photographs and front cover painting, 21.5cm x 14cm. Price £1.45.
ISBN 1-85279-079-2. - Reviewed by Paul D. Brock.
This book is part of a Your First series by TFH/Kingdom Books, which are designed
to provide clear, practical information, with full colour illustrations throughout; incredible
bargains at £1.45 each.
David Alderton will be known to many phasmid rearers for his Step by Step Guide to
Stick Insects, also published by TFH in 1992 - in fact, many of our members joined after
learning of the PSG from his book; they would probably otherwise have been unaware of our
existence. This new book contains similar sound advice on culturing species. The colour
photographs of several species do not, unfortunately, include Sipyloidea sipylus or the male
of Extatosoma tiaratum (both feature in Alderton’ s 1992 book). A strikingly beautifiil insect
is the gynandromorph (part male, part female) Oreophoetes peruana on page 17 - my
photograph of PSG member Gordon Ramel’s insect.
Any phasmid enthusiast will want a copy of this book, which should amply succeed in
its aim of helping the beginner. Buy it and wonder how the publishers can produce a book
at such an attractive price!
The Ecology of Java and Bali by Tony Whitten, Roehayat Emon Soeriaatmadja & Suraya
A. Afiff (1996), Published by Periplus Editions (HK) Ltd. 983 pages, plus 28 pages of
colour plates. 16cm x 23.5cm. Price £50.00. ISBN 962-593-072-8. - Reviewed by Phil Bragg.
This excellent book is volume two of The Ecology of Indonesia Series. In addition to
the colour plates, there are many black and white plates, drawings, maps and diagrams. The
book covers all aspects of the ecology of Java and Bali although the coverage of particular
groups of fauna, in chapter five, varies greatly because of difficulties obtaining reliable
information. This book will undoubtably provide an essential guide for anyone interested in
the ecology of the area. I make no attempt to review the book in detail, except for the
section on phasmids.
Stick insects and leaf insects are covered in more detail than most insect groups: on
pages 273-277, and there is one colour plate of Orxines macklottii at the end of the book.
The section on phasmids provides a usefol checklist of the 125 species recorded from these
two islands and indicates which species are endemic.
Atlas of grasshoppers, crickets and allied insects in Britain and Ireland by E.C.M. Haes
& P.T. Harding. (1997) The Stationery Office, London. 61 pages, 295mm x 210mm.
Price £15.50, ISBN 0-11-702117-2, -Reviewed by Phil Bragg.
This book gives distribution maps for all the native and naturalised orthopteroid insects
in Britain and Ireland. The phasmid section (pages 52-53) gives one map but does not
distinguish between the species; the distribution of the different species is discussed on the
facing page. The book has an attractive colour cover with photographs of six species of
Orthoptera. The price seems excessive for what is little more than an up-date of the maps
in Grasshoppers and allied insects of Great Britain and Ireland (Marshall & Haes, 1988).
Phasmid Studies, 6(2): 46
Phasmid abstracts
Video Review
Stick & Leaf Insects - A Novice’s Guide to Keeping Phasmids. GK Video, price £11.99.
- Reviewed by Paul D. Brock.
PSG members can benefit by ordering direct from GK Video, PO Box 213, Grimsby,
DN36 5ZG for a special mail order price of £10, inclusive of postage and packing; please
quote your PSG membership number when ordering. Foreign orders will be sent by surface
mail.
At last! A well filmed, practical video for the enthusiast, with some beautiful film
dealing with phasmid behaviour. This 32 minute film has a commentary by Keith Stiff, who
relays an incredible range of information.
I like the practical advice included in sections on moulting, housing, feeding, general
care and breeding. The feeding section even includes outdoor filming of obtaining bramble.
There is an excellent sequence dealing with the moulting of a male Extatosoma tiaratum. My
only criticism is that there are so many sequences of species early on that novices may be
confused as to which species is being shown, although some insects are named. However,
the ’star’ performers’ particulars are given in more detail at the end of the video.
The Phasmid Study Group is recommended and indeed the ’stars’ have been obtained
from members, with Roger Reeve and Peter Vice given particular mentions. This video is
a must for individuals and schools starting, or thinking of starting, cultures. PSG members
will wish to obtain a video even if they are experienced in breeding phasmids; it is worth it
just for the footage of Heteropteryx dilatata, Extatosoma tiaratum, Orxines macklottii, and
others.
Readers may be interested to know that this video is the second in a series: the first is
on tarantulas and the third (in production) will cover mantids.
Phasmid Abstracts
The following abstracts briefly summarise articles which have recently appeared in other
publications; also included are publications published since 1992 (the first issue of Phasmid
studies) which have only recently come to the attention of the editor. Some of these may be
available from local libraries. Others will be available in university or college libraries,
many of these libraries allow non-members to use their facilities for reference purposes free
of charge. In the UK libraries can usually obtain publications which they do not hold by
using the inter-library loan system; there is usually a charge for this service. A similar inter-
libraiy loan system operates in many other countries.
The editor of Phasmid Studies would welcome recent abstracts from authors so that they may
be included in forthcoming issues. In the case of publications specialising in phasmids,
Phasma and Le Monde des Phasmes, only the longer papers are summarised.
Arillo, A., Ortuno, V.M. & Nel, A. (1997) Description of an enigmatic insect from Baltic
amber. Bulletin de la Societe Entomologique de France, 102(1): 11-14.
In this paper an interesting orthopteroid-like insect preserved in Baltic amber is
described. The specimen has such peculiar features that the attribution to an order is
difficult. A comparison with other amber oithopteroid-like insects is also made.
Phasmid Studies, 6(2); 47
Phasmid abstracts
Bouchard, P., Hsiung, C.C. & Yaylayan, V.A. (1997) Chemical analysis of defense
secretions of Sipyloidea sipylus and their potential use as repellents against rats. Journal of
Chemical Ecology, 23(8): 2049-2057.
The defensive secretion of Sipyloidea sipylus was analyzed using a GC-MS technique.
Five chemicals were identified (diethyl ether, acetic acid, benzaldehyde, limonene, and
benzothiazole) and mixed in about the same ratio as detected in the secretion to determine the
potential effectiveness of such a mixture as a repellent against lab rats.
Brock, P.D. (1997) The jungle nymph, Heteropteryx dilatata (Parkinson). Bulletin of the
Amateur Entomologists’ Society, 56(412): 109-112, plates 97K & 97L.
Some general notes on Heteropteryx dilatata, with four colour photographs.
Brock, P.D. (1997) Magnificent Malaysian insects. Bulletin of the Amateur Entomologists’
Society, 56(413): 135-136, plates 97M & 97N.
The author briefly discusses various insects found on a trip to Sarawak and Peninsular
Malaysia. There are colour photographs of three phasmids: a female Lonchodes strumosus
(Brunner), a female Acacus sarawacus (Westwood), and a male of a new species of
Anarchodes from Fraser’s Hill, Peninsular Malaysia, The article includes photographs of
other insects and a general view of Fraser’s Hill.
Brock, P.D. (1997) Breeding the stick-insect Phasma gigas (L.) from Papua, New Guinea.
Bulletin of the Amateur Entomologists’ Society, 56(414): 181-184, plates 1-4.
Brief comments on rearing Phasma gigas, with some historical notes. Figure 1 shows
lateral and dorsal views of the egg [with labels transposed] . The plates are black and white
photographs of the adults.
Clark Sellick, J.T. (1997) Descriptive terminology of the phasmid egg capsule, with an
extended key to the phasmid genera based on egg structure. Systematic Entomology, 22:
97-122.
An attempt is made to standardize further the descriptive terminology of the phasmid
egg capsule by introducing stricter definitions and standard abbreviations. In addition, the
various forms of the internal micropylar plate are categorized. Eophasma Sellick is replaced
by Eophasmodes nov.n. A key to 131 generic forms of these eggs is provided. Where more
than one egg form is associated with a genus, a diagnosis of the subgroups is provided.
Fausto, A.M., Mazzini, M., Cecchettini, A. & Giorgi, F. (1997) The yolk sac in late
embryonic development of the stick insect Carausius morosus (Br.). Tissue & Cell, 29(3):
257-266.
Differentiation of the yolk sac was examined ultrastructurally and cytochemically in late
embryonic development of the stick insect Carausius morosus. During migration along the
yolk sac, endodermal cells form a discontinuous cell epithelium, leaving wide intercellular
channels between neighbouring cell clusters. Within the same cell cluster, cells are all joined
by septate junctions. In the proximity of the proctodeum region, intercellular channels are
filled with numerous cell debris which are shown to derive from vitellophages undergoing cell
lysis. Yolk sacs resolved by gel electrophoresis are shown to release a number of vitellin
polypeptides into the culture medium. These are equivalent in molecular weight to those
present in the vitellophage yolk granules. This observation is consistent with the evidence that
the basement lamina may act as a course physical filter, retaining particles larger than
colloidal thorium dioxide and allowing free percolation of peroxidase. Differentiating
Phasmid Susies, 6(2): 48
Phasmid abstracts
endcxiermal cells form a microvillar striated border along the apical plasma membrane. A
number of vesicular criptae were frequently seen in these differentiating endodermal cells.
Electron dense granules released by endodermal cells are suggested to play a role in
vitellophage lysis and vitellin release from the enclosed yolk granules.
Frantsevich, L. & Cruse, H, (1997) The stick insect, Obrimus asperrimus (Phasmida,
Bacillidae) walking on different surfaces. Journal of Insect Physiology, 43(5): 447-455.
Unrestrained adult stick insects {Obrimus asperrimus) walked below a tread wheel 4 or
30 mm wide in the upside-down position or above a ’bridge’ 30 or 60 cm wide in the upright
position. They were recorded on video and the positions of reference points on the legs and
on the body were measured on still frames. Step parameters such as step amplitude, step
duration, swing duration, body height and ground width are given for broad and narrow
footing as well as for leg trajectories and the course of leg joint angles. Joint angles were
calculated directly between the leg segment vectors. Walking with a narrow footing, the stick
insect used the same slow metachronal gait as on a broad footing. Adjustment to the narrow
footing was accomplished by narrowing the ground base, raising the body distance,
depressing the femora (by 40-45°) and flexing the tibiae (20-25° in the front and middle legs).
The upper turning-point of the swing movement seems to be determined by a constant
amplitude of the vertical movement component rather than a given position in a body-fixed
coordinate system. Walking on a horizontal rod of circular cross-section, the insects
preferred to walk upside down below the rod for small diameters, but preferred to walk
upright above the rod if its diameter was great enough.
[Editor’s note: The species used in this experiment belongs in the genus Aretaon Rehn &
Rehn, as Aretaon asperrimus (Redtenbacher)]
Gade, G., Lorenz, M.W. & Hoffmann, K.H. (1997) Stick insect (Carausius morosus;
Phasmatodea: Lonchodidae) brain extract contains multiple fractions with allatostatic activity.
European Journal of Entomology , 94(3): 361-368.
A first attempt to purify inhibitors of juvenile hormone biosynthesis from extracts of
brains of the Indian stick insect, Carausius morosus, is reported here. A heterologous
bioassay in the cricket, Gryllus bimaculatus, was used throughout the study. Separation of
a prepurified extract using C-18 reversed phase high performance liquid chromatography (RP-
HPLC) resulted in a broad zone with biological activity. Upon re-chromatography of each of
six active regions on a wide-pore C-18 column, the material separated in distinct peaks of
biologically active fractions. Introducing a third purification step (C-8 column) resulted in
a total of 18 absorbency peaks with allatostatic activity. It remains to be seen whether all
these fractions contain authentic allatostatins which are active in stick insects, or whether
other functions can be attributed to them.
Gorkom, J. van (1997) PSG 174 Lopaphus caesius (Redtenbacher). Phasma, 7(26): 18-20.
A report on rearing Lopaphus caesius from Vietnam. Measurements and colour
photographs are included for bodi the male and female; the egg is illustrated by a drawing.
Kittmann, R. (1997) Neural mechanisms of adaptive gain control in a joint control loop:
Muscle force and motoneuronal activity. Journal of Experimental Biology, 200(9): 1383-1402.
An adaptive gain control system of a proprioceptive feedback system, the femur-tibia
control loop, is investigated. It enables the joint control loop to work with a high gain but
it prevents instability oscillations. In the inactive stick insect, the realization of specific
changes in gain is described for tibial torque, for extensor tibiae muscle force and for
Phasmid Studies, 6 ( 2 ); 49
Phasmid abstracts
motoneuronal activity. In open-loop experiments, sinusoidal stimuli are applied to the
femoral chordotonal organ (fCO). Changes in gain that depend on fCO stimulus parameters
(such as amplitude, frequency and repetition rate), are investigated. Furthermore,
spontaneous and touch-induced changes in gain that resemble the behavioural state of the
animal are described. Changes in gain in motoneurones are always realised as changes in the
amplitude of modulation of their discharge frequency. Nevertheless, depending on the
stimulus situation, two different mechanisms underlie gain changes in motoneurones. (i)
Changes in gain can be based on changes in the strength of the sensorimotor pathways that
transmit stimulus-modulated information from the fCO to the motoneurones. (ii) Changes in
gain can be based on changes in the mean activity of a motoneurone by means of its spike
threshold: when, during the modulation, the discharge of a motoneurone is inhibited for part
of the stimulus cycle, then a change in mean activity subsequently causes a change in
modulation amplitude and gain. A new neuronal mechanism is described that helps to
compensate the low-pass filter characteristics of the muscles by an increased activation,
especially by a sharper distribution of spikes in the stimulus cycle at high fCO stimulus
frequencies. The work was done with Carausius morosus.
Marescalchi, O. & Scali, V. (1997) Chromosomal and NOR patterns in the polyclonal stick
insect Bacillus atticus atticus (Insecta; Phasmatodea). Genome, 40(2): 261-270.
Bacillus atticus atticus is a complex of thelytokous parthenogens, related to the bisexual
Bacillus grandii, that ranges from Sardinia to Near Astern countries. Karyotypic and
cytogenetic differentiation of the B. atticus atticus diploid unisexual "isolates" is really higher
than expected. Its standard karyotype has 2n = 34 chromosomes, but several instances of
repattemed or even aneuploid complements have been found. The number and location of
silver-stained NORs are particularly intriguing, since in addition to homozygous NOR
patterns, simple or double hemizygous strains are found spread over specific and wide
regions. The odd patterns are not due to Ag-NOR staining technique artifacts, since the FISH
method, using rDNA probes, apparently labels the same ribosomal clusters. Transpositions
and translocations have been suggested to account for some NOR patterns, but hybridizations
between different NOR-bearing races are also a possible cause. This chromosomal survey
clearly contributes to a better understanding of B. atticus phylogeny.
Miksys, S., Lange, A.B., Orchard, I. & Wong, V. (1997) Localization and neurohemal
release of FMRFamide-related peptides in the stick insect Carausius morosus. Peptides
(Tanytown), 18(1): 27-40.
FMRFamide-like immunoreactivity was localized immunohistochemically in the central
and stomatogastric nervous systems, visceral tissues, and the neurohaemal corpora cardiaca,
transverse, and segmental nerves. Each of these neurohaemal areas contains one
morphologically distinct type of immunoreactive neurosecretory granule. The haemolymph
level of FMRFamide-like peptides, quantified by RIA, is higher in animals sampled 2 h into
the dark cycle, relative to those sampled at mid-light cycle or 9 h into the dark cycle. High
potassium depolarization evokes the calcium-dependent release of FMRFamide-like peptides
from neurohaemal areas in vitro and HPLC fractionation of haemolymph, corpora cardiaca,
and their bathing medium suggests that these organs contribute a single peptide to the
FMRFamide-related peptides circulating in the haemolymph of active animals.
Potvin, J. (1997) Eieren van wandelende takken. Phasma, 7(26): 1-9.
Illustrations of the eggs of 104 species of phasmids; showing dorsal views only.
Phasmid Studies, 6 ( 2 ): 50
Phasmid abstracts
Scapigliati, G., Pecci, M., Piermateei, A. & Mazzini, M. (1997) Characterization of a
monoclonal antibody against a 180 kDa hemocyte polypeptide involved in cellular defence
reactions of the stick insect Bacillus rossius. Journal of Insect Physiology, 43(4): 345-353.
Defence properties of haemocytes were investigated using the anti-haemocyte
monoclonal antibody BrHl obtained by immunizing mice with 2% paraformaldehyde-fixed
haemocytes of the stick insect Bacillus rossius. In Western blot analysis, the antibody
recognized a ISOkDa antigen in haemocyte cell lysates, whereas fat body lysates and cell-free
haemolymph were negative. In hnmunofluorescence analysis of cultured or freshly collected
haemocytes, BrHl stained intracellular antigen(s) in detergent-treated cells. Transverse
cryosections of adult stick insects probed by immunofluorescence with BrHl showed in situ
the scattered distribution of haemocytes inside the haemocoel. The antigen(s) recognized by
BrHl appears to be involved in cell defence haemocyte-mediated mechanisms, as evidenced
by the fact that cryosections of insects challenged in vivo with yeast cells, bacteria, or
polystyrene latex particles and probed with BrHl showed an accumulation of antigen
surrounding the injected stimuli.
Sellick, J. (1997) The range of egg capsule morphology within the Phasmatodea and its
relevance to the taxonomy of the order. Italian Journal of Zoology, 64: 97-104.
Within the order Phasmatodea there is a basic egg capsule morphology which defines
the order. This shows extensive variation in detailed form. Some of this is abberation,
which can be recognised and excluded from taxonomic considerations. Much is adaptation
to egg-laying techniques and egg survival. Nevertheless the range of egg capsule form both
externally and internally is a valuable aid to the classification at sub-ordinal level and below
of this group of insects. In particular this confirms the Timematodea as members of the
order, confirms the distinct status of Phylliidae and shows some so far undivided groups such
as Necrosciidae to be taxonomically very diverse. A new tribe Extatosomatini is proposed
for Extatosoma Gray on the basis of both egg and adult morphology. A survey of internal
micropylar plate types within the order is given.
Viscuso, R., Narcisi, L., Sottile, L. & Giuffrida, A. (1997) Secretory product of the lateral
oviducts of Baculum thaii Haus. (Phasmida: Phasmatidae) and its change during egg transit.
International Journal of Insect Morphology and Embryology, 25(4): 369-379.
The secretory product elaborated by the epithelium of the lateral oviducts of Baculum
thaii (Phasmida: Phasmatidae) gives rise, in the oviduct lumen, to 2 main structural types:
fibrils and granules. The ultrastructural characteristics of the fibrils are uniform throughout
the various oviductal zones in both the virgin and mated females. However, the organization
of the granules is characteristic of a particular zone, and sometimes different in the same
zone, when virgins are compared with mated females. The fibrils accumulate on the chorion
of newly ovulated eggs to form a thick, compact sheath outside which the granules converge,
undergo rapid and multiple fragmentation, and overlay the fibrillar sheath in an orderly
arrangement. The fibrillar and granular components give rise, in both virgin and mated
females, to the formation of a sclerotized sheath called extrachorion that is still present in
newly laid eggs. The probable role played by these 2 extrachorionic components in relation
to the egg is discussed.
Zompro, O. (1996) Beitrage zur Kenntnis philippinischer Phasmiden II. Bemerkungen iiber
philippinische Obriminen, mit einer Neubeschreibung (Phasmatodea: Heteropterygidae:
Obriminae). Entomologische Zeitschrift, 106(11): 450-456. [In German]
A new genus and species of Phasmatodea, Sungaya inexpectata n.gen., n.sp. and its egg
Phasmid Studies. 6 ( 2 ): 51
Phasmid abstracts
are described from the Philippine island of Luzon. The form of the egg appears to be new
for the Obrimini. The male of Aretaon echinatus (StM, 1877) is described for the first time.
It' is similar to that of Aretaon asperrimus (Redtenbacher, 1906). Ecological notes on
Stenobrimus bolivari Redtenbacher, 1906 are included.
Publications noted
The following paper has been noted, but no abstract has been received.
Knutelski, S. & Royaud, A. (1997) Liparus dims Herbst (Coleoptera: Curculionidae) and
Clonopsis gallica (Charpentier) (Phasmatodea: Bacillidae), two interesting species for the
entomofauna of the Pyrenees- Atlantiques Department (France). Entomologiste, 53(3): 97-98.
An introduction to
Rearing Praying Mantids
A5 softback, 16 pages, 10 figs. ISBN 0-9531195-0-5.
This book describes methods of rearing and breeding praying mantids.
The headings include; An introduaion to praying mantids, Types of
mantids. Structure of mantids, Mantids in captivity. Cages, Feeding,
Breeding, Sexing, Mating, Egg laying. Identification, Preserving mantids.
Obtaining mantids. Distributing mantids, and Sources of further
information.
The book is illustrated with 10 black and white drawings, plus
one on the front cover. The drawings illustrate six different species of
mantids, how to distinguish the sexes, deuils of the fore leg, and an
internal view of an egg case.
An Introduction to
Rearing Cockroaches
by
PhUE. Bragg
An introduction to
Rearing Cockroaches
A5 softback, 16 pages, 14 figs. ISBN 0-9531195-1-3.
This book is .intended as a beginners’ guide to rearing cockroaches. It is
illustrated with 14 black and white drawings, plus one on the front cover.
The drawings illustrate eight different species of cockroaches and show
how to distinguish the sexes.
There is a general introduction to cockroaches with information
on the structure and different types. The commonly available species are
grouped according to general type and their suitability for culturing.
Cages, feeding, sexing and preserving are all discussed. There are
suggestions on obtaining and distributing cockroaches, and there is a list
of books offering further information.
Price: £2.50 each. Postage should be added at the following rates: UK 20p, Europe 70p, Elsewhere £1.10.
Orders and payment in pounds sterling should be sent to:
P.E. Bragg, 51 Longfield Lane, Ilkeston, Derbyshire, DE7 4DX, U.K.
Phasndd Studies, 6(2); 52
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