Ilja Svetnik, BSc - unipub

MASTER THESIS

Ilja Svetnik, BSc

RED LIST AND DNA BARCODING OF CARINTHIAN AND

STYRIAN CENTIPEDES (CHILOPODA)

For the master’s program

Ecology and Evolutionary Biology

UNIVERSITY OF GRAZ - 2019

ACKNOWLEDGMENTS

This work could not have been possible without the immense support of the University of Graz and the

outstanding Professors and Researchers of the Institute of Biology, as well as its head at the time Dr.

Christian Sturmbauer.

I want to thank my supervisor Dr. Stephan Koblmüller and my mentor Lukas Zangl, MSc, who supported

me in every way possible and shaped this thesis into what it has become. I also want to thank Dr. Werner

Holzinger for his immense help and advice in creating a cohesive Red List, and his support years ago,

when I just thought about maybe investing time in researching soil arthropods.

My thanks also extend to my previous supervisor Dr. Gernot Kunz, who supported me during my

bachelor thesis and thus laid the ground stone for this thesis’ research.

I also want to thank Dr. Nikolaus Szucsish from the Vienna Natural History Museum (NHM) for providing

me with additional material from different locations, thereby adding immensely to my barcoding data

bank.

Lastly, I want to thank all the people who I had contact to and who offered their support during the

making of this thesis, including but not limited to Dr. Karin Voigtländer, Dr. Christian Wieser, and Dr.

Henrik Enghoff as well as my colleagues Michaela Bodner, David Fröhlich and Maximilian Wagner.

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ABSTRACT

Centipedes are predatory, multi-legged arthropods inhabiting soil, litter, and rocky surfaces in all kinds of

habitats. As such they serve an important role in the global detritus chain and soil ecology. Central

European centipedes have been studied ever since the 18th century with a focus on alpine species and

species from the British Isles. Austria has been a hotspot of these studies ever since, and has been

known to be home to around 70 native species of centipedes. Unfortunately, these species have been

described multiple times with different names and have been plotted together and split apart

taxonomically various times in the last decades, leaving taxonomists confused, especially due to the lack

of consistent literature. The scarcity of taxonomic information and difficulty of evaluating population

sizes leads to a lack of red lists on centipedes, especially those with reasonable criteria. DNA barcoding is

a recent method that can assist in studying the phylogeny of any living being that is gaining popularity,

yet should be used with great care. Today many ecologists turn to analysis of DNA or RNA in order to

study the diversity of beings. We take advantage of both classic and new methods in order to resolve the

taxonomy and status of native centipede species.

TABLE OF CONTENTS

ACKNOWLEDGMENTS ................................................................................................................................... 2

ABSTRACT ................................................................................................................................................... 3

TABLE OF CONTENTS ..................................................................................................................................... 4

INTRODUCTION ............................................................................................................................................ 6

Concerning Centipedes 6

Evolutionary and taxonomic history 6

Taxonomy and morphology 7

Anatomy and physiology 9

Mating and reproduction 10

Systematics and species numbers 11

Distribution 12

Prey and predators 13

Venomousness and danger 14

Concerning Barcoding 16

The “Barcode of Life” Projects 17

Part I.

Red List of Centipedes 19

State of the art 19

Methodology 22

Disclaimer 23

Results 25

Checklist of Carinthia 25

Checklist of Styria 26

Statistical analysis 27

Legend of the Red List 28

Red List of Centipedes of Carinthia 29

Red List of Centipedes of Styria 41

Critical reflection 52

Summary and outlook 54

Part II.

DNA Barcoding of Centipedes 55

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State of the art 55

Methodology 58

Results and discussion 60

Summary and outlook 64

APPENDIX ................................................................................................................................................. 68

BIBLIOGRAPHY ........................................................................................................................................... 69

INTRODUCTION

Concerning Centipedes

Centipedes can be generally described as predatory arthropods of the taxonomic class Chilopoda, part of

the subphylum Myriapoda, which also includes the Millipedes (Diplopoda) and two other multi-legged

arthropod groups, the Pauropods and Symphylans (ROSENBERG 2009). According to MINELLI (2011) there

are 3461 species of Centipedes described, while ADIS & HARVEY (2000) claim, there are at least 5000

species out there that have yet to be described. These numbers make the Centipedes a relatively small

group within the enormous phylum of arthropods, and yet they are quite diverse, show a wide variety of

adaptations, and are present in almost every habitat on every continent except Antarctica. They can all

be described as worm-like, multi-legged, chitinized inhabitants of the ground and litter-layers that vary in

size from a few millimeters up to more than 30 centimeters in length (ROSENBERG 2009). Despite their

name, no Centipede species has exactly one hundred legs, since all species show an odd number of leg

pairs. The number of individual legs varies from 30 to 354 depending on the genus. Inside the class of

myriapods, the Centipedes are an exclusively carnivorous clade of fast, rather strong, venomous

predators (LEWIS 2011).

Evolutionary and taxonomic history

The myriapods in general are a very old taxonomic group, with the earliest records of specimens dating

back to the late Silurian, 430 million years ago (SHEAR 1992). It is generally accepted today that the

insects and myriapods (together referred to as “Antennata”) made their way from sea to land two

separate times in evolutionary history. Convergent selection processes led to the myriapod ancestors

losing the first pair of antennae and slowly evolving a homonomic segmentation of their bodies. Thanks

to a strong drive of evolutionary forces at the end of the Silurian, a wide variety of prehistoric myriapod

species could already be found in the early Devonian. The records of these prehistoric myriapods also

present some of the largest species of land-dwelling Arthropods we know of. The genus Arthropleura

brought forward enormous, heavily armored specimens with a length of up to 2.3 meters (BRADDY et al.

2008). The heavily armored Arthropleura armata (MEYER 1854) were probably more closely related to the

Millipedes than Centipedes, but it has long been suggested by many scientists that they could have been

predators, a thesis that lacks evidence until this day (SCOTT; CHALONDER & PATERSON 1985). Records of

prehistoric Centipede species are present, but none of them represent enormous species. Predatory

Centipedes ever since filled the economic niche of ground- and litter-living macro predators and have

ever since been of greater importance in the detritus food-chain. Today, Centipedes are omnipresent in

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almost all habitats in the world. Depictions of Centipede species can be found everywhere in recorded

history, especially in sacred texts of ancient Egypt and medical records of ancient Greek society, with the

earliest depictions dating back to our Cro-Magnon ancestors and their cave-paintings (LEWINSON &

LEWINSON 2004). It was Greek philosopher ARISTOTLE who first described Centipedes around 300 B.C.,

merely characterizing them as “wingless insects with several legs”. Hundreds of years later, in 1758,

French naturalist Carl LINNÉ used a similar description to distinguish between the genera Iulus and

Scolopendra, which LINNÉ both placed inside the order of the Aptera, or wingless insects (VERHOEFF 1928).

The genera Iulus and Scolopendra later turned into the taxonomic groups Diplopoda and Chilopoda we

use today. Modern zoologists like VERHOEFF, KOCH and ATTEMS dedicated themselves so intensively study

the myriapods, and thereby pioneered the descriptions and pre-genetic systematic order of these

animals, leading the way for today’s researchers to approach the order with modern scientific tools and

challenge the discoveries made decades ago.

Taxonomy and morphology

DNA-based research by EDGECOMBE & GIRIBERT (2002) has shown us, that the evolutionary gap between

the myriapod orders Diplopoda (Millipedes), Pauropoda and Symphyla is quite small, while there is a

huge gap between those orders and the Centipedes (Chilopoda). Together, the Diplopoda, Pauropoda

and Symphyla make up the taxon Progoneata, named after the shared position of the genital system

inside the animals’ bodies. The second characteristic these groups share, is their plant-, fungi- or

microbial-based diet (HOPKIN & READ 1992). The carnivorous Centipedes therefore have a different body

structure, one that is slicker, less stiff and makes them much faster and maneuverable.

EDGECOMB & GIRIBERT (2009) primarily distinguish the centipedes by their head-structure and branching of

the stigmata. We differentiate the Notostigmomorpha, who have short, single stigmata on the posterior

area of each dorsal plate, from the Pleurostigmomorpha, who feature deep-reaching, heavily branched

stigmata that are always present in pairs. The head structure of the Notostigmomorpha is rounded, and

(unlike all other myriapods) they feature compound eyes, and rather long, multi-segmented antennae.

This taxon only consists of one order: the Scutigeromorpha, also known as house centipedes, which

consist of 3 families, 2 of them only native to tropical areas. Only the family Scutigeridae is present in

Europe, with only one species: Scutigera coleoptrata LINNÉ 1758, thanks to its high abundance in human

populated areas also referred to as Common House Centipede. Within the Pleurostigmomorpha we find

all other taxa of Centipedes, with the main orders Lithobiomorpha (stone centipedes),

Craterostigmomorpha, Geophilomorpha (soil centipedes) and Scolopendromorpha (tropical centipedes).

There is very little known about the order Craterostigmomorpha, which is only represented by two

species found in Tasmania and New Zealand, and therefor will not be further discussed.

The Scutigeromorpha, along with the above mentioned Craterostigmomorpha, and the yet to be

discussed Lithobiomorpha make up the subphylum (or division) called Anamorpha. The group comprises

the centipedes whose number of body-segments increases as the animals mature, with the young

hatching with only 7 pairs of legs and the adults always having 15 pairs. In contrast, the young specimens

of the Epimorpha group (consisting of the Geophilomorpha and Scolopendromorpha) all hatch with their

full number of leg-pairs, which for the Scolopendromorpha is 21 or 23 (39/43 in the case of

Scolopendropsis duplicate EDGECOMB & GIRIBERT 2008) and anywhere between 25 and 177 for the

Geophilomorpha (SIRIWUT et al. 2016). Both the Lithobiomorpha, Craterostigmomorpha and

Geophilomorpha have a heteronomic body-segmentation, but while all body segments bear legs within

the stone centipedes, only every second segment bears legs within the soil centipedes. Only the

Scolopendromorpha show a homonomic body-segmentation, with every segment bearing a pair of legs.

The Scutigeromorpha have 15 segments, covered by seven tergites.

The house centipedes have compound eyes and long antennae composed of only two segments. All

other species of centipede feature a single Ocellus, a group of them, or don’t feature eyes at all. They

also feature multi segmented, (mostly) short antennae, which can be moved in all directions and serve as

sensory appendages. The heads of all centipedes are flat (except for the Scutigeromorpha), and feature

three pairs of mandibles, the latter two in the form of maxillae. The appendages of the first body-

segment have evolved into maxillipeds, in the form of strong pincers with venom glands, which serve

both to paralyze and catch prey as well as to defend against enemies. Except for the last pair of

appendages, all other extremities serve as legs. Like all other arthropods, centipede legs are made up of

several parts, and consist of a coxa, trochanter, femur, tibia and (depending on the genus) a monomial or

binomial tarsus, always equipped with a claw-like pretarsus. The last pair of legs usually serves a

different purpose and is always shaped differently from the other appendages. They usually serve as

either grabbing- or sensory extremities. The last body-segment only features a very small pair of

extremities, the gonopods, positioned close to the genital opening and used to deliver the

spermatophores or the placement of eggs. Only the Scolopendromorpha have no visible gonopods,

which are shaped like telescopes, and can therefore be retreated inside of the pregenital body-segment.

The gonopods are very diverse in shape, and therefore often used to distinguish even closely related

species. Especially the gonopods of the females are much longer and larger. They consist of three

segments, are mostly claw-shaped and often feature spurs. The gonopods of the males are much shorter

and stubbier and cover the extendable penis (ROSENBERG 2009).

One of the most interesting (and most discussed) structures of the centipede body are the coxal pores,

which are present (at least) on the last pair of extremities in almost every species in some shape or form.

Inside the Lithobiomorpha the pores are always present on the coxae of the last four leg-pairs, mostly in

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low numbers, as a single row of elongated wells. Only few genera like Eupolybothrus feature smaller,

oval pores in much higher number, distributes scree-like on the coxae. We find similar patterns of coxal

pores within the orders Geophilomorpha and Scolopendromorpha, where the pores are not always as

easy to recognize as such. The functions and evolution of said pores has been a hot topic of discussion

ever since, and even though a multitude of studies tried to pin down its reason of existence, evidence is

sparse and often holey. Factually, the exact functions and biological meaning of said pores are yet to be

fully discovered, but today it is generally accepted, that the pores have some function in the water-

regulation of the centipedes, and serve as openings to control the intake and release of water vapor.

They might additionally release fragrances or secretes, but (most definitely) do not function as excretory

organs to release venoms, sticky secretes or slime to cover their bodies or eggs, as assumed by science in

previous decades (ROSENBERG 2009). To fully discover the functions and biological meaning of the coxal

pores, additional research and the application of new methods will be necessary. But as new are

constantly being developed and the field of science turning towards the analysis of arachnid- and insect-

secretes, we may answer this question sooner than we think.

Anatomy and physiology

The inner anatomy of centipedes is the same we find in all arthropods. They have a hemolymph which is

distributed inside the worm-like body by multiple ostia and the heart tube. A ventral nerve cord (NVC)

links the many extremities to a tritocerebrum (supraesophageal ganglion), thereby ensuring a well-

coordinated motor function. Excretion happens through a malpighian tube system, branching tubules

lined by chitin extending from the alimentary canal, found in all higher arthropods. Uric acid is produced

and released through the anal opening, water and ions are being reabsorbed at the rectum, a key

adaptation of land-dwelling arthropods, especially the thin-skinned species and those living in arid areas

such as deserts. As a fact, specimens of the Geophilomorpha lose water at a much slower rate than the

Lithobiomorpha, which may be contributed to the pleural membrane of the specimens, which shows

much higher levels of sclerotization. The size and shape of the stigmata, as well as the ability to curl, also

have an impact on the water regulation of the animals. Questionable, again, is the role of the coxal pores

in this instance. Primal myriapods also feature one or two pairs of plesiomorphic glands at the antennae

and maxillary nephridia, of the same kind found in the closely related Pancrustaceans. The exchange of

air functions just as it does within the class of insects, through trachea, which deliver oxygen directly to

the muscles (RUPPERT, FOX & BARNES 2004). As discussed earlier, the shape and position of the stigmata,

the openings of the trachea, vary from order to order and are therefore a taxonomic trait.

Mating and reproduction

Centipedes are ovipary creatures. The reproductive process happens without a real mating or

copulation. Instead the males place their spermatophores on a weave and leave it there for the females

to pick it up. A real pairing only happens within the Scutigeromrpha, where the males attract the females

with a dance, place their spermatophore on the ground once a female shows up, and in the process pull

the dancing female over the spermatophore, sticking it onto the body of the female. Very few species of

centipedes are also known to be parthenogenetic and produce offspring out of unfertilized eggs. All

other female specimens of centipedes pick up the spermatophores of the males and fertilize the eggs

inside of their bodies themselves. They then lay a small to moderate amount of eggs, ranging from ten to

50 within the Scolopendromorpha and Geophilomorpha, and even less within the Lithobiomorpha. The

Scutigeromorpha usually also lay between ten and 50 eggs, but within laboratories they have shown to

produce anywhere from 60 to 150 eggs, depending on the conditions. The egg deposition occurs in

temperate areas in spring and summer, with some species occurring earlier than others, and some

species even having two breeding events within one season. Parental care is not unusual within the

centipedes. The Geophilomorpha and Scolopendromorpha (as well as the Craterostigmomorpha)

females guard their eggs and young until several days after they hatch. The females curl around their

offspring in a protective state, with the Geophilomorpha curling around them with the dorsal side, with

the defensive secretory glands positioned to the outside, while the Scolopendromorpha (and

Craterostigmomorpha) curl around them with the ventral side, defending their offspring with the

(mostly) heavily chitinized dorsal plates. The timespan between the deposition of the eggs and the

hatching of the offspring is very variable from species to species and can take anywhere between one to

several months. Where parental care is happening, the mother will constantly lick her eggs and keep

them clean from any spores of fungi. Once the offspring hatches the young are quite inactive for a

couple of days and will only leave the protection of their mother after a couple of moultings at a more

active state. In the meantime, the young depend on their mother, and are being fed by her indirectly.

There are few species of Scolopendromorpha which are known to always be matriphagous, where the

body of the mother serves as food for her offspring. When the mother is disturbed by a predator, she

will either abandon her offspring or consume them. Abandoned eggs will quickly be destroyed by fungi

without the constant care of their mother. There is no evidence of any parental care within the

Scutigeromorpha and Lithobiomorpha. The females will place their eggs separate and either mask them

with litter material or place them inside of individual little holes in the ground (LEWIS 2007). Hatched

“larvae” of centipedes do look very similar to the adult individuals but are mostly transparent. Those of

the Lithobiomorpha, Craterostigmomorpha and Scutigeromorpha have fewer legs when they hatch, and

grow two or more pairs of legs with each moulting until they reach the full number of lags. The

Scutigeromorpha are born with only four pairs of legs, most Lithobiomorpha with seven and the

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Craterostigmomorpha with 12. They usually reach their full number of leg pairs before they are fully

adult, since the last two to three moultings of the young serve to develop the gonopods, additional

antenna segments, different glands and some ocelli. The Gophilomorpha and Scolopendromorpha hatch

with the same number of legs they carry as an adult, and only develop the gonopods and additional

glands during each moulting until they are adults (LEWIS 2007). Usually centipedes reach sexual maturity

at an age of one year and are fully adult after one to three years. The time between the hatching of the

eggs and full adulthood can vary a lot from species to species and ecozone to ecozone, especially with

the larger species of Scolopendromorpha, who can grow to reach an age of multiple years. The average

lifespan of a centipede is somewhere between four to seven years, with Scolopendra subspinipes (LEACH

1816) reaching an age of over 10 years, according to YATES III. & JULIAN (1992). Since centipedes lay a

moderate amount of eggs, have quite long incubation times and need several years to become fully

adult, it is commonly accepted that they target K-selection, and do not target massive distribution of

their species, but rather seek to preserve a rather high number of individuals over multiple generations

(ALBERT 1979).

Systematics and species numbers

The Lithobiomorpha, or stone centipedes consist of two similar families, the Henicopidae and

Lithobiidae. The Henicopidae consists of 19 genera and approximately 120 species, while the Lithobiidae

consists of over 80 genera and around 1200 described species worldwide. The Scolopendromorpha, or

tropical centipedes, comprises five families. Even though the name suggests otherwise, the

Scolopendromorpha are not exclusively found in the tropics, but all over the world. The family

Mimopidae is the smallest, consisting of only two valid species of the order Mimops. The family

Plutoniumidae has only recently been established. There are now six species described, either from the

new genus Plutonium or the older taxon Theatops, both formerly contributed to the family Cryptopidae.

The family Cryptopidae is still quite large, and consists of almost 200 species, of which over 150 can be

contributed to the genus Cryptops. The family Scolocryptopidae consists of around 100 species, with the

two large genera Newportia (over 50 species) and Scolocryptops (20 species) contributing majorly. Lastly,

the family Scolopendridae consists of several hundred species. This family is where we find the largest

centipede species in the world, especially from the genus Scolopendra (around 100 species), which is also

where we find the only two amphibious centipedes we know of, Scolopendra cataracta SIRIWUT,

EDGECOMBE & PANHA 2016 and Scolopendra paradoxa DOMÉNECH 2018 (HOLMES 2016, BATES 2016). The

Geophilomorpha, also called soil centipedes, are another very diverse group, consisting of at least seven

families and almost 1300 species, making them the most diverse group of centipedes (MINELLI 2011). The

family Geophilidae (including the former Linotaneidae, Dignathodontidae and Macronicophilidae) alone

features over 120 genera. The remaining families Mecistocephalidae, Oryidae, Himantariidae,

Schendylidae, Zelanophilidae and Gonibergmantidae are all significantly smaller, but do add to the

extreme diversity found in soil centipedes (MYERS et al. 2013). Soil centipedes lack eyes, just like all

Cryptops and most Scolocryptops and Theatops species. Because they spend most of their life in the

ground or dark caves, they also sometimes lack pigmentation, or show reduced pigmentation and

chitinization in most areas of their bodies, save for the heads.

Distribution

Thanks to their far-reaching evolutionary history and prehistoric dominance, the centipedes can be

found all over the world, in almost every climate zone and altitude, with many species adapting to

extreme conditions such as extreme heat, high atmospheric pressure and regular flood events. Most

centipedes prefer wet, lush layers of litter as their habitat, while the tropical species live on and below

the massive vegetation of the rainforests, in caves as well as deserts, coastlines and even completely

underground. The larger species of the rainforests and desert areas usually have individual dens and lairs

inside of wood and vegetation or under rocks and larger systems of boulders (LEWIS 2007). Those species

native to central Europe also occur under rocks and dead wood but are scarce outside of these habitats

and rarely ever found on vegetation. Geophilomorpha usually live underground or inside the uppermost

layers of the ground, but there are several species that live in caves or inhabit rock systems, and a

handful of examples of species that dwell on coastlines made up from sand, gravel or boulders (LEWIS

1961). Those species occurring in higher altitudes usually show higher levels of pigmentation, a

phenomenon that can often be observed within the arthropods. There are no records of centipede

species in the snow zones of mountains or the Arctic regions, but there are species who live beyond the

arctic circle (RIEGER ET AL. 2010). Centipedes are generally sensitive to light and extreme heat but can

comprehend extreme heat much better than extreme cold. Centipedes occur in deserts, steppe areas

and savannas much more frequently than in the tundra, and there are no species present in any ice-

deserts (LEWIS 2007, ROSENBERG 2009). Those species who live in countries with cold winters like Austria,

survive the cold by retreating underground, inside protected microhabitats or spend the winter inside

dead wood or large boulders. Those centipedes that are more robust and thermophilic can also be found

in cities or man-made landscapes. These are mostly very common species with few preferences and

demands when it comes to their habitat. In central Europe these are species such as Lithobius forficatus

(LINNÉ 1758) and Scutigera coleoptrata (LINNÉ 1758), in tropical areas also the larger Scolopendra

specimens (LEWIS 2006). The thermophilic Scutigera coleoptrata (LINNÉ 1758) is often found in or near

houses in rural regions or submontane settlements and doesn’t seem to be bound to any specific

habitat. Scutigera are very active and fast hunters and can often be found under matrices, carpets and

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pillows (RICKS 2015, LEWIS 2007). In contrast to Scutigera and some millipedes (i.e. Ommatoiulus

sabulosus (LINNÉ 1758)), Lithobiomorpha and Geophilomorpha are usually not found in houses, maybe

because they are bound to litter- and ground habitats. Native species of the Scolopendromorpha do also

appear in cities, gardens and parks, but often do not stay there for very long, unlike Scutigera does.

Especially the genus Cryptops finds at least two species in flowerbeds of large cities (even with neozoic

species like Cryptops rucneri MATIC 1967) and even Scolopendra cingulata LATREILLE 1829 can be found

breeding in gardens in the northeastern parts of our country.

Prey and predators

A key element for the presence or absence of centipedes is the abundance of possible prey and natural

enemies in a certain habitat (LEWIS 2007). Centipedes have no problem with the presence of conspecifics

or other species of centipedes in the same are, if there is enough prey for them to survive. They are less

frequent in areas where hunting-spiders and larger species of ants are present, but do live side by side

with millipedes, since they do not share a common prey but prefer similar habitats. While millipedes,

such as the Julidae, feed on fungi and dead woods, they produce perfect microhabitats for smaller

arthropods such as springtails (Collembola), which serve as a primary prey for smaller species of

centipedes, who therefore prefer to live in the very same microhabitat as its prey. They are predatory

generalists and hunt a multitude of different animals. They never consume plants but have shown to

feed on some dead plant material or fungi when brought close to starvation. They usually hunt and

consume all animals that are rather thin-skinned and not much larger than the centipedes themselves

(LEWIS 2007). The hypo- and endogenous Geophilomorpha mostly feed on smaller and larger species of

worms like Nematodes or smaller Oligochaeta, which are omnipresent in soil and easy to hunt and kill

(WEIL 1985). Thanks to their sizes, the giant centipedes within the order Scolopendromorpha are even

capable of hunting smaller vertebrates such as mice, small lizards and amphibians. They can catch bats in

Venezuelan caves out of the air by crawling onto the ceiling of the cave and letting the posterior parts of

their serpent-like bodies hang down. If they manage to grab a flying bat, they let themselves fall to the

ground and immediately paralyze the prey with a powerful bite and the injected venom (MOLINARI et al.

2005). Giant tropical centipedes such as Scolopendra gigantea (LINNÉ 1758) have even been reported to

feed on serpents, birds and bats, and there are even reports of snakes consuming entire Scolopendra

specimens alive which then proceeded to consume the snake themselves from the inside, eventually

hatching out of their predators’ body. Springtails and other small ground-arthropods serve as the main

prey for most small and mid-sized species of centipedes, but they are also intraguild predators, and are

known to hunt other centipedes, spiders and other competitors. While little is known about the

predatory behavior of the Scutigeromorpha and Craterostigmomorpha, we can assume that they are

similar to those of the Lithobiomorpha and Scolopendromorpha (LEWIS 2007).

Even though the most centipedes live a very cryptic life, they do have to face a lot of predators. They

themselves are hunted by mice, lizards and snakes, as well as birds and larger beetles. Interestingly

enough, these are the same groups of animals which serve as prey to the largest species of centipedes.

Larger centipedes are also a common prey of Mongoose. We know of two species of animals who

exclusively feed on centipedes: the African ant Stigmatomma1 pluto (GOTWALD & LEVIEUX 1972), feeding

exclusively on a species of Geophilidae, and the south-African snake Aparallactus capensis SMITH 1849

(LEWIS 2007). It has been observed time and time again that centipedes avoid the neighborhood of ants,

especially larger, colonial ants, probably because of the high aggressivity and precaution of those

species, who will attack and kill any potential danger to the colony. This claim is further supported by the

fact that millipedes can often be seen fairly close to colonial ants. The slow, xylobiontic millipedes aren’t

of any danger to the colony and might therefore be accepted as neighbors, unlike their fast, predatory

centipede relatives.

Venomousness and danger

Centipedes use their massive maxillipeds to kill their prey and defend against enemies. The maxillipeds

are formed as massive claws, which always feature glands filled with venomous substances, which

means that all centipede species are venomous, except for a few cryptic species which do not produce

venomous substances or even lack the glands to do so. In addition to these glands, the ventral plates of

the Geophilomorpha also feature microscopic glands which produce sticky secretes with venomous

components such as Benzoic acid and hydrogen cyanide. We assume that the Lithobiomorpha are also

able to produce such sticky secretes, but many descriptions of such glands date back to the time when

the above mentioned coxal pores were described as secretory glands, which they are certainly not

(EISNER T., EISNER M. & SIEGLER M. 2005). The biggest advantage of the centipedes is their immense

flexibility and speed. When being exposed to light, hearing an approaching predator or being disturbed,

they can retreat into the smallest spaces between rocks or underground in no time.

In the field of zoology, centipedes are often an overlooked group of arthropods, positioned somewhere

between the insects and arachnids, and mentioned sparsely next to venomous spiders and scorpions.

Despite that, centipedes have been known to be venomous predators just as long as spiders and

scorpions, with some species of the Scolopendra order even being of great danger to human beings

(KNOFLACH 2010). In ancient Egypt and Mesopotamia, we can find countless sculptures and depictions of

1 formerly genus Amblyopone.

https://en.wikipedia.org/w/index.php?title=William_H._Gotwald,_Jr.&action=edit&redlink=1

15

centipedes in temples and medical reports, next to the classic images of serpents and snakes. The

ancient Egyptians even had a god by the name of Sepa resemble a centipede, who was known as a

protector against evil animals, and described as a god of death, disease and healing. NEWPORT (1844) was

the first scientist to describe the maxilliped claws of the centipedes, even though they have already been

depicted a few decades earlier in the zoological drawings of LEEUWENHOEK. The venom glands evolved

from an aggregate of epidermal glands along the cuticular canal. The glands are holocrine, and venom is

released through contraction of the muscles surrounding the glands (KNOFLACH 2010). How potent the

venom of the centipedes really is, has been a topic of discussion for decades. While most centipedes use

their venom to kill or paralyze worms, springtails and snails, larger specimens of the Scolopendromorpha

are even capable of paralyzing small vertebrates like mice, frogs, lizards, fish and bats (LEWIS 1981;

CARPENTER & GILLINGHAM 1984). The venom of Scolopendra species has undergone a lot of research in the

last decade. COHEN & QUISTAD (1998) were some of the first scientists to discover the key component of

the centipede venom. Those components are histamine, serotonin and different enzymes, which form a

complex protein mix that has both neuro- and cytotoxic attributes (GOMES et al. 1982; MOHAMED et al.

1983). In 2006 WENHUA et al. were able to isolate an unknown peptide from the venom of Scolopendra

subspinipes (LEACH 1816), given the name Scolopendrin I., and shown to have immobilizing or even

destructive effects on a wide variety of different microorganism. In 2009 GONZALES-MORALES et al.

detected the first phospholipase in the venom of Scolopendra viridis SAY 1821, and studies by RATES et al.

from 2007 revealed that at least 12 components of the Scolopendra venom have shown to be highly

toxic to most insects. A controversial topic has ever since been the effect of the Scolopendra venom on

humans. Reports of cases of death caused by centipede venom date back to the first century A.D. but are

met by modern scientists with much disbelief (BÜCHERL 1946, 1971; MINELLI 1978; LEWIS 1981). Only one

incident reported by PINEDA (1923) is to be believed. The reported death of a small child in the

Philippines caused by a bite of Scolopendra subspinipes (LEACH 1816) is accepted as fact by multiple

authors (REMINGTON 1950; BUSH et al. 2001) who examined the case. SERINKEN et al. (2005) report the

death of a man from Turkey who died after being bitten by a centipede, but explain that his death was

actually caused by an infection after the bite, not the venom or the bite itself. All other modern reports

of centipede bites do not suggest any lethal effects on mankind at all, not even the bites of the

enormous Scolopendra gigantea LATREILLE 1829 (REMINGTON 1950; SOUTHCOTT 1978; MUMCOUGLU &

LEIBOVICI 1989; MOHRI et al. 1991; RODRIGUEZ-ACOSTA et al. 2000; BARROSO et al. 2001: BUSH et al. 2001;

BOUCHARD et al. 2004; GUERRERO 2007; VARIANZADEH et al. 2007; IVOCHEVA et al. 2008). Bites of larger species

of centipedes are reported to be extremely painful and have a long-lasting burning sensation. They also

provide a large risk of dangerous inflammation and infections (KNOFLACH 2010). Bites of smaller species

are often compared to bites of ants and usually only hurt for a minute without leaving any scars for

inflammations behind. Reports of bites from Geophilomorpha are sparse, but there are several species

whose bites are known to be quite painful, often compared to the sting of a bee or wasp.

Scutigeromorpha bites are also not reported a lot, which seems odd, since they are the most abundant

species of centipedes in houses, and are known to be the most “aggressive” centipedes, or rather the

most likely to bite in defense, and release a rather large amount of venom, which results in a very

unpleasant bite, though the pain only last for a short amount of time. In general, unlike spiders and

snakes, centipedes use their venom very sparingly, and only in extreme emergencies. They are regarded

as fairly aggressive animals and use their venom mainly to paralyze and kill their prey. It is assumed that

centipedes are capable of biting without releasing any venom, which further supports the claim that

centipede venom is extremely precious (ASPÖCK 2010). Most native species of centipedes are too small to

even penetrate human skin and are therefore regarded as completely harmless. Only bites of Scutigera

coleoptrata (LINNÉ 1758) and Geophilus electricus (LINNÉ 1758) are unpleasant, with the latter species

barely coming in contact with humans.

Concerning Barcoding

DNA barcoding is a novel system designed to provide rapid, accurate, and automatable species

identifications by using short, standardized gene regions as internal species tags (HEBERT & GREGORY

2005). VALENTINI, POMPANON & TABERLET (2008) described the rise of barcoding as a tool for ecologists as a

new method that has received much attention, and will increasingly be used to not only identify a single

species from a specimen or its remains, but also determine the species composition of environmental

samples. Ecologists can even take advantage of DNA tools when only hair, feces or urine left behind by

animals are available for species identification. As a consequence, it will make the Linnaean taxonomic

system more accessible, with benefits to ecologists, conservationists, and the diversity of agencies

charged with the control of pests, invasive species, and food safety. VALENTINI et al. (2008) further claim

that “short DNA fragments persist in the environment and might allow an assessment of local

biodiversity from soil or water”. The ideal DNA barcoding marker has not yet been found or may not

even exist, because the marker would have to meet a wide variety of criteria:

(i) The gene region sequenced should be nearly identical among individuals of the same

species, but different between species.

(ii) It should be standardized, with the same DNA region used for different taxonomic groups.

(iii) The target DNA region should contain enough phylogenetic information to easily assign

unknown or not yet ‘barcoded’ species to their taxonomic group (genus, family, etc.).

(iv) It should be extremely robust, with highly conserved priming sites and highly reliable DNA

amplifications and sequencing (particularly important when using environmental samples,

containing a mixture of many species to be identified at the same time).

17

(v) The target DNA region should be short enough to allow amplification of degraded DNA.

Usually, DNA regions longer than 150 bp are difficult to amplify from degraded DNA.

The authors also note that, different groups of users will give different priorities to those categories: “for

taxonomists, criteria (i), (ii) and (iii) will be more important […], whereas ecologists working with

environmental samples will favor criteria (iv) and (v) […]”. The approach most taxonomists take is

barcoding sensu stricto, whereas ecologists favor a sensu lato approach. The now well established

initiative CBOL (consortium for the barcode of life) has been designed to fit with DNA sequencers based

on capillary electrophoresis, which yield a typical read length of 500–1000 bp. CBOL proposes the 658 bp

gene region encoding the mitochondrial cytochrome c oxidase 1 (COI) for standardized barcoding of

animals. Yet the very characteristics that make the COI gene a candidate for high throughput DNA-

barcoding, also limits its information content on a deeper phylogenetic level, as HEBERT et al. (2003) point

out. According to MORITZ & CICERO (2004) DNA barcoding should therefore not be misinterpreted as a

tool to resolve the “tree of life”. More and more sequence data for the accepted barcoding markers are

becoming available in public databases as sequencing facilities improve. VALENTINI et al. (2008) note: “[…]

the quality of the sequence data in GenBank, EMBL or DDBJ is not always perfect, either as a result of

sequencing errors, contaminations, sample misidentification or taxonomic problems”. CBOLs own

database specially dedicated to DNA barcoding called BOLD (Barcode of Life Data Systems) provides an

efficient and accurate way for species identification. Not only does the database record DNA sequences

of several individuals per species, but also contains complete taxonomic information, place and date of

collection, and specimen images.

The “Barcode of Life” Projects2

ABOL – the Austrian Barcode of Life, is an initiative that aims to generate a public database of DNA-

barcodes of all native lifeforms, by building a national network of researchers in the field of biodiversity.

Thereby the platform additionally promotes all national biodiversity research and coordinates related

projects all over the country. It has been designed as a long-term initiative of at least 10 years and began

its run in 2017, after a 3-year pilot phase from 2014 to 2017. Initiated and coordinated by the Museum

of Natural History in Vienna, its pilot phase and coordination is being financed by the Austrian Federal

Ministry of Education, Science and Research, and involves all relevant institutions involved in biodiversity

research, such as universities, museums, biosphere parks and several different research institutions in

addition to federal conservation departments and regional scientific societies. The initiative consists of

group-specific clusters, characterized by a group of experts of each specific group of lifeforms or specific

2 Source: www.abol.ac.at

orders of lifeforms. Inside the ABOL project is a specific cluster of myriapods and basal hexapods,

coordinated by Dr. Nesrine Akkari and Mag. Daniela Bartel of the Museum of Natural History in Vienna.

The coordination team of ABOL also aims to stay in contact with other national barcode-of-life-initiatives

such as iBOL, GBOL and CROBOL. Like all other BOL-initiatives, ABOL stands for quality standards. The

specimens from which the DNA is extracted have been legally collected and are deposited in a publicly

accessible scientific collection. DNA-barcodes and corresponding required meta-data are fed into the

ABLO-database. All work from the collection to the documentation, sampling and evaluation process are

carried out obeying highest carefulness and meeting laboratory standards. Remaining DNA or tissue

samples are being stored in a collection, and all barcodes are incorporated into the international BOLD©

database.

19

PART I.

RED LIST OF CENTIPEDES

State of the art

Native centipedes have been studied ever since the days of Austrian entomologist Robert LATZEL, who in

1880 published his monography on the myriapod-fauna of the then Austro-Hungarian empire. LATZEL was

the first Austrian scientist to build upon the works of pioneers such as Pierre André LATRILLE, who earlier

published works on the fauna of Austria, Hungary and parts of Czechoslovakia. LATZELS work, with

contributions by naturalists Carl Ludwig KOCH and Frederik Vilhelm August MEINERT who described

several new species native to Austria, thereby filled a huge gap in European faunistic research. LATZEL

took KOCHS and MEINERTS descriptions and documentations and combined them into one collective work,

which to this day is regarded as the most important collection of descriptions for native myriapods. The

collection consists of 130 taxa of Diplopoda (1 genus, 2 subgenera, 69 species and 56 forms) and over 40

groups of Chilopoda (2 genera, 29 species and 12 forms), as well as 4 taxa of both the Pauropoda and

Symphyla. In 1884 LATZEL published his second work on the Pauropoda, Symphyla and Diplopoda,

another work of huge importance in the field of myriapod systematics and distribution. LATZEL also

turned out to be a pioneer in the field of gonopod-research and was the first to realize that size and

shape of the gonopods are the best and often only way to determine certain species of myriapods. LATZEL

thereby majorly contributed not only to the field of Austrian faunistics, but also to the field of

entomology globally. His work was so far reaching, there have only been a dozen new native species of

centipedes described since LATZELS days. LATZELS work has been described by other entomologists as

extremely detailed and highly accurate, and his collection of myriapods is regarded as one of the most

important collections of the museum of natural history in Vienna, containing several type-species.

LATZELS impact on the field of myriapod-research will remain unforgotten, thanks to a group of

researchers who gave a group of house-centipedes the taxonomic name Latzelia. LATZEL himself always

knew there were more species to be discovered, and wanted other naturalists to become “friends” of

the myriapod-group. Only then, as LATZEL himself wrote: “[…] can we expect to achieve completion”.

LATZEL himself only published two more works in the following years, both on the myriapod-fauna of

Carinthia, and worked on the index of myriapods of Tyrol with HELLER and DELLA TORRE.

The next naturalist to contribute to the Austrian myriapod-research was German zoologist Karl Wilhelm

VERHOEFF, who worked as a lecturer at the University of Bonn in Germany at the time, where he

previously graduated. He traveled through most of the southern lands like Istria, Dalmatia and Carniola,

and visited parts of Tyrol and Styria. With his works VERHOEFF mainly aimed to describe more and more

species of myriapods, and in the process learned a lot about the affinity of certain species. At the same

time, Austrian monarch and zoologist Carl August of Attems-Petzenstein (a.i. Carl ATTEMS) began his

private studies on Styrian myriapods. ATTEMS wanted to explore the fauna of Styria, and previously

studied the gonopods of Polydesmida. In 1895 he published a revised work on the fauna of Styria, which

also featured numerous publications and findings from Lower Austria. With VERHOEFFS help, ATTEMS

described over 1800 new species of myriapods, which he collected on numerous private excursions all

around the world and published 138 scientific papers on the topic. In the following years more and more

publications described new and known species of Polydesmida native to Austria and Germany, all

building upon the foundations laid by ATTEMS, who later included said descriptions in his work on the

systematics of Styrian Polydesmida (HÖLDER 1901).

Edward Holt EASON was the next naturalist to describe several new species of myriapods in the 1960’s,

mainly species of the British Isles. He published an extraordinarily accurate determination key for

centipedes, including some of the best zoological drawings of centipede bodies and gonopods to date.

Several excellent works on the biodiversity of Austrian centipedes existed by the 1960’s, what was

missing is a uniform collection, to combine all the scattered descriptions and information that would also

clarify synonymic species, unknown species and establish a uniform taxonomy. We got that with the

infamous “Catalogus Faunae Austriae”, published in 1972 by naturalist Marcus WÜRMLI, who included a

uniform determination key like the one of EASON.

Another notable work was published in the early 1990’s by biology-professor and zoologist Armin KOREN,

who collected several species of centipedes in Carinthia and published two determination keys for the

native species, which included drawings by EASON and a large amount of new drawings by KOREN himself.

On the basis of his own publications, KOREN later with the nature conservation union and local

government, worked on Carinthias red list of centipedes, the very first red list for centipedes of Austria.

Unfortunately, the list was not very detailed, and left out questionable species and other key elements

like the responsibility for the species and abundance of the species in neighboring countries and states.

With the rise of phylogenetic methods in the last decade, the field of systematic zoology changed its

face. The systematic system that has been valid for centuries was changed from its foundations, thanks

to discoveries from genetic studies, which by now have mostly replaced the classic morphological

approach to systematics. This also impacted the field of entomology and thereby the systematics of

myriapods. Today, more and more new species are being found and described every year, while

21

phylogenetic studies indicate that different previously described species are either one and the same or

represent either sub-species or variates. This “enlightenment” led to a fundamental problem, which is

(again) the establishment of a uniform and valid systematic system. For many years this has gotten more

difficult, especially since many scientists today rely on genetic data only, even if its results heavily

contradict morphological discoveries. The growing number and lack of definition of sub-species further

complicates the establishment of a uniform system. A system that combines morphological and

phylogenetic evidence is a difficult task many entomologists face.

Since it is almost impossible to keep a record of any larger centipede population (or its area of

distribution), and the fact that this has not been done in the last 25 or more years, there is no red list of

Centipedes released under the umbrella of the IUCN. The IUCN has only evaluated ten species of

Centipedes as of January 2020, all of them of tropical origin, mostly from the Seychelle Islands. There are

checklists and regional red lists of centipedes from several German states (SPELDA 1999, 2004, 2005;

VOIGTLÄNDER 2004, 2005; REIP & VOIGTLÄNDER 2009; DECKER & HANNING 2011), as well as one for the country

of Germany based on said lists (GRUTTKE et al. 2016). Most of these lists work with data from findings and

only give abundances and responsibilities. Same goes for the only red list of centipedes in Austria from

1990, where KOREN lists the native centipedes by abundance and habitat preference to evaluate the

endangerment of at least those species with enough records.

The Austrian soil conservation law has been part of the nature conservation decree ever since its

initiation (BGBl. Nr. 491/1984), but is directed by the nine states, not the nation. The importance of its

quality and faunal diversity is often underestimated. Today we have a much clearer vision of the

biodiversity of cryptic taxa such as Collembola, Acari and Diplura, but where the taxonomy becomes

clearer every day, very little is known about the distribution and abundance of these groups of animals.

The idea behind creating a full checklist and red list of Carinthian and Styrian centipedes is to raise

awareness of the biodiversity of the taxa, importance of some of its species (especially endemic species)

and create a better understanding of the biology, ecology and role of centipedes in the environment.

Some species have the potential to be used as indicators in environmental studies or nature

conservation surveys.

Future friends of these taxa can improve upon this work and aim to one day evaluate population status

and distribution of these species throughout Austria. Only then can we work on a cohesive red list of

centipedes following IUCN criteria.

Methodology

The south of Austria is the most studied part of the country when it comes to centipedes, and therefore

shows the highest density of records and descriptions of said animals. According to RABITSCH & ESSL

(2009) the level of records of Carinthia and Styria is almost on par with those of the “British Myriapod

Survey”, yet the rest of the country is far behind. There are around 70 species of centipedes present on

Austrian soil, though statements regarding that number may vary. Pinning down the exact number of

native species is much more challenging than it would seem to be. With the rise of internet databases

such as Fauna Europeae (ENGHOFF 2007) and ChiloBase (MINELLI 2006) many species underwent

taxonomic revision. Species such as Geophilus noricus VERHOEFF 1928 and Geophilus pauropus ATTEMS

1927, which only a decade earlier had just been distinguished and morphologically described as separate

by KOREN, completely disappeared and are ever since synonymous with a third species, Geophilus

oligopus (ATTEMS 1895). Many of the native subspecies described by WÜRMLI as endemic met the same

fate. There are some species described as native that lack proper description or records, while others are

heavily implied to be native but have not been found yet. The exact species count an author will give

heavily depends on whether or not he will count said species as native or not, and if the author agrees

with all the synonyms made by ENGHOFF, MINELLI and others in the early 2000’s.

For this thesis, the number of valid described species for Austria can be set somewhere between 60 and

70. Of these species between 50 and 60 can be found in the counties of Carinthia and Styria. The species

count of the two neighboring counties is almost the same, and either one could practically have the

higher amount of species. For the county of Carinthia, we set the number at 52 certain species and an

additional 2 species with unclear status. For Styria, the count is 51 certain species with an additional 2

uncertain species. The “certainty” of said species of course is never 100%, leaving at least some leeway

as is always the case in taxonomy. With 59 described species and possibly even 60, the two counties

could be home to over 90% of all centipede species native to Austria.

A quantitative comparison and statistical analysis based on the number of individuals in a certain area is

practically impossible when it comes to centipedes. A solid basis for any analysis is the number of

discovery sites. In addition to the number of sites the bond of a certain species to a specific habitat

should be considered for evaluations. For stenoecious species this proves to be particularly difficult, as

very little is known about the preferences of most of these animals (GRUTTKE et al. 2016). Especially with

these “rare” species, another problem arises. An unconfirmed or missing entry of a discovery site may

have a large impact on any statistical evaluation of species with few localities, which (in reality) does not

impact the overall trend of the population.

23

Due to the low degree of discovery, it is impossible to formulate any trend criteria for a population. As

centipedes are k-strategists, population fluctuations are common, often extreme, and almost

unpredictable (GRUTTKE et al.. 2016).

Disclaimer

The following chapter contains the two red lists for the counties of Carinthia and Styria respectively. For

the county of Carinthia, part of the neighboring Eastern Tyrol was included for all statistical analysis.

Other authors including KOREN (1986, 1992) did the same thing, because Carinthia and Eastern Tyrol

share the same ecogeological and meteorological area.

In order to create a cohesive and logical checklist of native centipedes, research was done using the

currently valid taxonomy used on the website Chilobase© (MINELLI 2016), distribution data of native

centipedes found on the website Zobodat (REICHL 1972) and the works of KOREN (1986, 1992), and

additional information from the website FaunaEuropeae©. The author did not concern himself with the

subject of subspecies, but comments regarding important subspecies have been added to the red list

separately. One of the major problems creating any checklist of species are synonyms. Centipedes have a

long history of countless revisions and name changes. The most encountered synonyms of the native

species have been added to the list, while a complete list of synonyms can be found in the appendices.

The states of Carinthia and Styria have been divided into units of squares by drawing lines between the

latitudes and longitudes. A square was “occupied” whenever data of a species’ presence between its

borders was available. To formulate statements on the abundance of a species, the total number of

occupied spaces (squares containing data) per state was taken as reference (Figure 1, p. 27). A 5-step

logarithmic scale was formulated to determine a species’ distribution from tiny scale (1) to omnipresent

(5). For category (5) the number of occupied squares necessary was set at 80, for category (1) at least 2.

Species with data from only one square were put in category (0), a category for species with deficient

data.

The two counties have been divided into 3 geographical subareas, all covering approximately the same

number of squares for further analysis of distribution. These areas were used as additional indicators of

the species’ distribution. If a species is present in all three areas, its category will be upgraded by one (for

example going from category (2) to (3) with less than 15 occupied squares), if a species is only present in

one area its category will be downgraded by one (going from category (3) to (2) despite occupying over

15 squares). There is no downgrading a species from categories (1) to category (0), as only species with

less than 2 records will be put in the latter. There is also no upgrading for species with more than 80

occupied squares that are present in three areas. Said species will feature a (!) icon next to their

category, indicating the impossibility of an up- or downgrade.

25

Results

Checklist of Carinthia

Eupolybothrus grossipes (C. L. KOCH, 1874)

Eupolybothrus tridentinus (FANZAGO, 1874)

Harpolithobius anodus (LATZEL, 1880)

Lithobius aeruginosus* C. L. KOCH, 1862

Lithobius agilis C. L. KOCH, 1874

Lithobius austriacus* (VERHOEFF, 1937)

Lithobius borealis MEINERT, 1868

Lithobius burzenlandicus VERHOEFF, 1931

Lithobius castaneus NEWPORT, 1844^

Lithobius crassipes* C. L. KOCH, 1862

Lithobius denatus C. L. KOCH, 1844

Lithobius erythrocephalus C. L. KOCH, 1847

Lithobius forficatus (LINNÉ, 1758)

Lithobius lapidicola MEINERT, 1872

Lithobius latro MEINERT, 1872

Lithobius lucifugus C. L. KOCH, 1862

Lithobius macilentus C. L. KOCH, 1862

Lithobius melanops NEWPORT, 1845

Lithobius microps MEINERT, 1868

Lithobius moellensis VERHOEFF, 1940 ?!

Lithobius mutabilis C. L. KOCH, 1862

Lithobius muticus C. L. KOCH, 1847

Lithobius nodulipes LATZEL, 1880

Lithobius pelidnus HAASE, 1880

Lithobius piceus C. L. KOCH, 1862

Lithobius punctulatus C. L. KOCH, 1847 ?

Lithobius pygameus LATZEL, 1880

Lithobius subtilis LATZEL, 1880~

Lithobius tenebrosus MEINERT, 1872

Lithobius tricuspis MEINERT, 1872

Lithobius validus MEINERT, 1872

Lamyctes emarginatus (NEWPORT, 1844) ~

Eurygeophilus pinguis (BRÖLEMANN, 1896)

Clinopodes flavidus C. L. KOCH, 1846

Stenotaenia linearis (C. L. KOCH, 1835)

Geophilus alpinus MEINERT, 1869

Geophilus electricus (LINNÉ, 1758)

Geophilus flavus (DE GEER, 1778)

Geophilus oligopus (ATTEMS, 1895)

Geophilus pygmaeus LATZEL, 1880

Henia illyrica (MEINERT, 1870)

Henia vesuviana (NEWPORT, 1845)

Strigamia acuminata (LEACH, 1815)

Strigamia crassipes (C. L. KOCH, 1835)

Strigamia transsilvanica (VERHOEFF, 1928)

Pachymerium ferrugineum (C. L. KOCH, 1835)

Schendyla carniolensis VERHOEFF, 1902

Schendyla nemorensis (C. L. KOCH, 1837)

Schendyla tyrolensis (MEINERT, 1870)

Dicellophilus carniolensis (C. L. KOCH, 1847)

Cryptops hortensis (DONOVAN, 1810)

Cryptops parisi BRÖLEMANN, 1921

Cryptops rucneri MATIC, 1967

Scutigera coleoptrata (LINNÉ, 1758)

52 confirmed species (54 total)

*Subgenus Monotarsobius (previously Genus) ~ not yet recorded, most definitely native ? Taxonomy uncertain ?! Taxonomy uncertain, possibly endemic ! endemic to Austria !! endemic to the state

Checklist of Styria

Eupolybothrus fasciatus (NEWPORT, 1845)




Lithobius aeruginosus* C. L. KOCH, 1862

Lithobius agilis (C. L. KOCH 1874)

Lithobius anisanus VERHOEFF, 1937

Lithobius austriacus* (VERHOEFF, 1937)



Lithobius castaneus NEWPORT, 1844 ~

Lithobius crassipes* C. L. KOCH, 1862

Lithobius curtipes C. L. KOCH 1847

Lithobius denatus C. L. KOCH, 1844


Lithobius forficatus (LINNÉ, 1758)

Lithobius franzi ATTEMS, 1949 !!





Lithobius macrocentrus ATTEMS, 1949 !


Lithobius microps MEINERT, 1868 ~




Lithobius peggauensis VERHOEFF, 1937 ?!

Lithobius pelidnus HAASE, 1880^


Lithobius pygameus LATZEL, 1880




Lamyctes emarginatus (NEWPORT, 1844)




Geophilus electricus (LINNÉ, 1758)














Scutigera coleoptrata (LINNÉ, 1758)

51 confirmed species (53 total)

*Subgenus Monotarsobius (previously Genus) ~ not yet recorded, most definitely native ? Taxonomy uncertain ?! Taxonomy uncertain, possibly endemic ! endemic to Austria !! endemic to the state

Statistical analysis

DATA:

Carinthia 9.536 km² Area 1 Area 2 Area 3 Eastern Tyrol 2.020 km² 3 360 squares 123 109 128 Styria 16.401 km² 544 squares 180 203 161

27957 km² 904 squares OCCUPIED SQUARES

4 Carinthia 189 47 47 63 Styria 163 63 72 26

352 STUDIED AREA IN % Carinthia 52,5 Styria 30,3

39

SPECIES NO. OF EACH RL CATEGORY

Cat. CAR STY CAR % STY %

0 6 10 11,5 19,6

1 5 7 9,6 13,7

2 6 3 11,5 5,9

3 17 13 32,7 25,5

4 13 10 25,0 19,6

5 5 8 9,6 15,7

Sum 52 51 100 100

3 not the entire area of Eastern Tyrol was considered for analysis, only the easternmost parts, approximately 1/3 of

the actual area. 4 Geographical subunits containing data of findings within its borders (Graphic 1).

Graphic 1: Areas with collection data in both Carinthia (left) and Styria (right).

Legend of the Red List

RED LIST CATEGORIES

Table 1: Categories and criteria of the red list

0 1 2 3 4 5

Occupied squares = 1 ≥ 2 ≥ 4 ≥ 15 ≥ 40 ≥ 80

Distribution category - tiny scale small scale normal spacious omnipresent

Abundancy category

Data deficient very rare rare moderate common very common

ADDITIONAL SYMBOLS

↑ = Category upgraded (species present in 3 areas),

↓ = category downgraded (species present in only 1 area)

! = category should be down-/upgraded, but not possible (species is already in Cat. 1 or 5)

ECOLOGY CATEGORIES

Temp = preferred temperature, Hum = preferred humidity, Alt = preferred altitude, Soil = preferred soil

acidity, Hab = preferred habitat

BOND CATEGORIES

? = questionable, ! = strong, 0 = low/none, !! = very strong5

1 2 3

Temperature cold neutral warm

Humidity low normal high

Altitude colline/sub mountain mountain/subalpine alpine/nival

Soil alkine none/either or acidic

1 2 3

Habitat open land, gardens, anthropologic areas

litter, rocks, wood, mixed habitats

deciduous forests, pine forests, mixed forests

5 Only used in Habitat category.

29

Red List of Centipedes of Carinthia


Synonyms: Polybothrus dubius MANFREDI,1948; Lithobius montanus C.L. KOCH, 1847; Lithobius grossipes debilis LATZEL,1889


Synonyms: Lithobius leptopus (LATZEL,1880); Polybothrus cerberus (VERHOEFF,1929); Polybothrus leptopus ssp. (VERHOEFF,1935)


Synonyms: Harpolithobius calcivagus VERHOEFF, 1925; Harpolithobius andreevi MATIC & STAVROPOULOS, 1988

Lithobius aeruginosus (C. L. KOCH, 1862)

Synonyms: Monotarsobius aeruginosus (VERHOEFF 1937)

Comment: The species is often listed

under its old genus name

Leptopolybothrus. LATZEL (1880) listed the

species under the name Eupolybothrus

leptopus.

Comment: KOREN names the subspecies

anodus and denatus, and distinguishes

them by the size of their gonopods H.

anodus denatus MATIC, 1957 is recorded

in Romania and “Yugoslawia”. H. anodus

anodus LATZEL, 1880 the species native to

Austria. The records from Carinthia may

be relict populations from the last glacial

period.


Synonyms: Lithobius paradisiacus MATIC & DARABANTU, 1971

Lithobius austriacus VERHOEFF, 1937

Synonyms: Monotarsobius austriacus (VERHOEFF, 1937)


Synonyms: Lithobius lusitanus würmanus (VERHOEFF 1937); Lithobius lapidicola (LATZEL 1880)


Synonyms: Lithobius gridellii MANFREDI, 1955; Monotarsobius veronensis MATIC & DARABANTU,1971

Lithobius castaneus NEWPORT, 1844

Synonyms: Lithobius lucasi NEWPORT, 1849; Lithobius meridionalis FEDRIZZI, 1877; Monotarsobius remyi VERHOEFF,1943

Comment: Very closely related to

Lithobius aeruginosus.

Comment: KOREN claims that the two

subspecies burzenlandicus and euxinicus

are recorded in Romania, while the

subspecies carinthicus is native to

Carinthia. Findings from Slovenia only

name the species with no subspecies

(KOS 1988). The findings of KOREN

generally match those made by MATIC in

1966. The valid name for the native

species is L. burzenlandicus carinthicus

nov. ssp. and can only be found in

Carinthia.

Comment: Records of this species only

exist from several areas in Carinthia, but it

is most definitely also native to Styria and

possibly even Burgenland. The species is

very circummediterrian and is distributed

throughout northern Italy, Slovenia and

western Hungary. According to MINELLI

(1987) the species inhabits temperate

oak-beech forests.

31

Lithobius crassipes C. L. KOCH, 1862

Synonyms: Lithobius (Monotasobius) crassipes C. L. KOCH, 1862; Lithobius atrifrons SILVESTRI, 1896, Lithobius podokes ATTEMS, 1903

Lithobius dentatus C. L. KOCH, 1844

Synonyms: Lithobius alpestris LATZEL, 1880


Synonyms: Lithobius armatus SSELIWANOFF, 1880; Lithobius dubius TÖMÖSVÁRY, 1880; Lithobius dubius TÖMÖSVÁRY, 1880

Lithobius forficatus (LINNÈ, 1758)

Synonyms: Lithobius leachii NEWPORT 1844; Lithobius vulgaris LEACH, 1817


Synonyms: Lithobius pusillus (LATZEL 1880); Lithobius sulcatus (C. L. KOCH 1862)

Comment: Originally described as L.

dubius by TÖMÖSVARY, in 1880, L.

erythrocephalus has been described by

many authors all throughout Europe. In

many places this species is distinguished

into two sympatric subspecies. The

subspecies L. erythrocephalus schuleri

(MATIC, 1966 schulleri) described by

VERHOEFF (1925, 1937) is much more

common in Austria, but the exact

taxonomy of the species and its

subspecies is in need of revision.

Comment: The “common stone centipede”

is the most common species of centipedes

all throughout Europe and is omnipresent

in almost any habitat.


Synonyms: Lithobius mutabilis transalpinus LATZEL, 1880


Synonyms: Lithobius alpinus C. L. KOCH, 1862; Lithobius rupivagus VERHOEFF, 1937; Lithobius walachicus VERHOEFF, 1901


Synonyms: Lithobius aulacopus LATZEL, 1880


Synonyms: Lithobius glabratus C.L. KOCH, 1847; Lithobius lusitanus crissolensis VERHOEFF, 1935

Lithobius microps MEINERT, 1868

Synonyms: Lithobius duboscqui BRÖLEMANN, 1896; Lithobius olivarum VERHOEFF, 1925; Sigibius puritanus CHAMBERLIN,1913

Comment: Though many sources cite

microps as missing in Austria, the species

has been recorded in Vienna, Lower

Austria and Carinthia, and is possibly

present in more states, since it is native to

most surrounding countries such as

Germany, the Czech Republic, Slovakia,

Hungary, Slovenia, Croatia, Albania, Italy

and Switzerland.

33

Lithobius moellensis VERHOEFF, 1940

Synonyms: none


Synonyms: Lithobius daday TÖMÖSVÁRY, 1880; Lithobius communis C.L. KOCH, 1844; Lithobius maculatus MATIC & DARABANTU, 1971


Synonyms: Lithobius bicolor TÖMÖSVÁRY, 1879; Lithobius cinnamomueus L. KOCH, 1862


Synonyms: Lithobius athesinus VERHOEFF, 1937


Synonyms: none

Comment: L. moellensis is the possible

third endemic species of centipedes in

Austria, and the only species endemic to

Carinthia. It has only been found and

recorded once, but the specimen is

damaged and has only been recorded to

have been found “in the Glockner area,

some 1500 meters above sea level”. The

taxonomy of the species is uncertain, and

even though it is listed as its own species,

has not officially gained endemic status

yet.

Comment: VERHOEFF noticed the variety of

certain traits of this speices and described

several subspecies: L. p. allemanicus from

Schwarzwald, L. p. insubricus from Wallis,

L. p. ponalensis from Gardasee and L. p.

annulipes from Hallstadt. A new

subspecies L. p. triangulatus nov. ssp. has

been described by KOREN and found at

the Saualpe.


Synonyms: Lithobius marginatus FEDRIZZI, 1877; Lithobius fanzagoi FEDRIZZI, 1876

Lithobius punctulatus C. L. KOCH, 1847

Synonyms: Lithobius matici (PRUNESCU1966)

Lithobius pygmaeus LATZEL, 1880

Synonyms: Lithobius rucneri MATIC, 1966


Synonyms: Lithobius nigrifrons LATZEL & HAASE, 1880; Lithobius balcanicus MATIC, 1973


Synonyms: Lithobius dolomiticus ATTEMS, 1903; Lithobius oligoporus LATZEL, 1885

Comment: Not to be confused with the old

taxonomic name for L. validus. The

taxonomy of this species is completely

uncertain and highly doubtful. There are

two records of L. punctulatus from the

Carinthian border in Eastern Tyrol.

35


Synonyms: Lithobius leptotarsis MATIC, 1959; Lithobius molleri VERHOEFF, 1893; Lithobius punctulatus machadoi PRUNESCU, 1966

Eurygeophilus pinguis (BRÖLEMANN, 1869)

Synonyms: Chalandea scheerpeltzi ATTEMS,1952; Geophilus baldensis VERHOEFF,1901


Synonyms: Geophilus montanus (MEINERT,1870); Geophilus gorizensis (LATZEL,1880)


Synonyms: Geophilus brevicornis C.L. KOCH,1837; Himantarium caldarium MEINERT,1886; Geophilus ormanyensis ATTEMS,1903


Synonyms: Geophilus insculptus ATTEMS, 1895; Geophilus glacialis VERHOEFF, 1928; Geophilus glocknerensis VERHOEFF, 1940

Comment: Its old name Chalandela

scheerpeltzi (ATTEMS 1952) is still used

but not valid anymore. This species is

native to the Pyrenees mountains, the

Cottian alps and the isle of Corsica. The

species has been recorded in seven

different areas in Carinthia. These may be

relict populations from before the last ice

age, which means the species could also

be found in other Austrian counties.

ChiloBase© (2016) also cites findings

from Slovenia, Switzerland, Italy and

Britain.

Comment: According to VERHOEFF (1938)

the genus Clinopodes (ATTEMS 1929)

should be replaced by the genus

Geophilus, but Clinopodes remains as a

valid taxon. There are three subspecies

described: the “new” subspecies C. fl.

carinthicus LATZEL is synonymous with the

subspecies C. fl. flavidus ATTEMS. The

subspecies C. fl. styriacus ATTEMS 1895

and C. fl. polytrichus ATTEMS 1903 are

also native to Austria.

Comment: The old genus name

Clinopodes is being used by KOREN and

EASON but is invalid.

Comment: Old invalid names include

Geophilus linearis, Geophilus

glocknerensis, Geophilus proximus,

Geophilus glacialis, Geophilus anglicanus

and Geophilus alpinus.

Geophilus electricus (LINNÈ, 1758)

Synonyms: Geophilus sudeticus HAASE, 1880; Geophilus helveticus VERHOEFF, 1928


Synonyms: Necroploeophagus longicornis EASON 1960; Geophilus longicornis LEACH, 1815; G. pygmaeus styricus VERHOEFF, 1895


Synonyms: Geophilus noricus VERHOEFF, 1928; Geophilus pauropus ATTEMS, 1927; Geophilus insculptus ATTEMS, 1895


Synonyms: Geophilus carnicus VERHOEFF, 1928; Geophilus cispadanus SILVESTRI, 1896; Geophilus larii VERHOEFF, 1934

Henia illyrica (MEINERT, 1870)

Synonyms: Chaetechelyne herzegowinensis VERHOEFF,1938; Henia termena CHAMBERLIN, 1952

Comment: Necroploephagus longicornis,

Geophilus longicornis, Geophilus

hortensis and Pachymerium tristanicum

are all old taxonomic names that are still

used but invalid.

Comment: Synonymous with the species

G. noricus (VERHOEFF 1928 and G.

pauropus (ATTEMS 1927), which have

been described as separate species by

KOREN and others.

Comment: According to ATTEMS 1929, G.

pygmaeus is an Illyric species native to

Slovenia, Croatia and Hungary. One

record from Carinthia and one from Styria

confirm the species is native to Austria. It

is also heavily implied that the species is

native to Upper Austria as well.

37

Henia vesuviana (NEWPORT, 1845)

Synonyms: Chaetechelyne vesuviana NEWPORT, 1845; Chaetechelyne corsica VERHOEFF,1943; Scolopendra fusca FOURCROY,1785


Synonyms: Scolioplanes italicus VERHOEFF,1928; Linotaenia rosulans C.L. KOCH, 1847; Geophilus subtilis C.L. KOCH, 1838


Synonyms: Geophilus breviceps NEWPORT,1845; Scolioplanes mediterraneus VERHOEFF,1928


Synonyms: Scolioplanes transsilvanica (VERHOEFF, 1935)


Synonyms: Geophilus caucasicus ATTEMS,1903; Geophilus paradoxus TÖMÖSVÁRY,1880; Mecistocephalus punctilabium NEWPORT,184


Chaetechelyne is valid no more. It’s a

Mediterranean species and native to

northern Africa, south France, Croatia and

all throughout Italy to Tyrol, Romania and

Hungary. The few records of this species

from Carinthia are probably relict

populations of warmer ages.


Synonyms: Schendyla (Echinoschendyla) zonalis BRÖLEMANN & RIBAUT, 1911


Synonyms: Dicellophilus nemorensis C. L. KOCH, 1837; Geophilus gracilis HARGER,1872


Synonyms: Schendyla montana ATTEMS, 1895; Brachyschendyla montana RIBAUT & BROLEMANN,1927


Synonyms: Geophilus apfelbecki VERHOEFF, 1898; Geophilus austriacus MEINERT, 1886; Mecistocephalus hungaricus TÖMÖSVÁRY,1880


Synonyms: Cryptops aenarienis VERHOEFF, 1943

Comment: The subgenus is

Echinoschendyla (BRÖLEMANN & RIBAUT),

but the old name Echinoschendyla zonalis

is invalid.


Dicellophilus is invalid.

Comment: The names Brachyschendyla/

Schendyla montana are not valid

anymore.

39


Synonyms: none

Cryptops rucneri MATIC, 1967

Synonyms: none

Scutigera coleoptrata (LINNÈ, 1758)

Synonyms: Scutigera pretzmanni WÜRMLI, 1973; Scutigera rubrovittata VERHOEFF, 1905

Comment: Closely related to Cryptops

hortensis, and maybe even synonymous.

The identity of Cryptops rucneri is

uncertain and only very few data exists on

the species.

Comment: This species is the only repre-

senttative of the Scutigeromorpha in

Europe and very thermophile. It has

invaded almost every anthropomorphic

area thanks to carryover but has no high

abundances. It is extremely anthropo-

genic and common in houses and

manmade structures. There are only few

populations in the wild, which is why the

species is often listed as endangered.

KOREN (1992) and Zobodat© (2016) show

no records of Scutigera in Carinthia and

Styria, yet it is omnipresent. It is

discussed if the species should be

classified as a Neozoon, but experts claim

the species populated Austria from the

south by itself (ESSL &RABITSCH 2002).

1 2

3

4

5 6

41

Red List of Centipedes of Styria

Eupolybothrus fasciatus (NEWPORT, 1845)

Synonyms: Lithobius montellicus (FANZAGO, 1874); Polybothrus praecursor alarichi (ATTEMS, 1934)


Synonyms: Polybothrus dubius MANFREDI,1948; Lithobius montanus C.L. KOCH, 1847; Lithobius grossipes debilis LATZEL,1889


Synonyms: Lithobius leptopus (LATZEL,1880); Polybothrus cerberus (VERHOEFF,1929); Polybothrus leptopus ssp. (VERHOEFF,1935)


Synonyms: Harpolithobius calcivagus VERHOEFF, 1925; Harpolithobius andreevi MATIC & STAVROPOULOS, 1988

Comment: The species is often listed

under its old genus name

Leptopolybothrus. LATZEL (1880) listed the

species under the name Eupolybothrus

leptopus.

Comment: KOREN names the subspecies

anodus and denatus, and distinguishes

them by the size of their gonopods H.

anodus denatus MATIC, 1957 is recorded

in Romania and “Yugoslawia”. H. anodus

anodus LATZEL, 1880 the species native to

Austria. The records from Carinthia may

be relict populations from the last glacial

period.

Lithobius aeruginosus (C. L. KOCH, 1862)

Synonyms: Monotarsobius aeruginosus (VERHOEFF 1937)


Synonyms: Lithobius paradisiacus MATIC & DARABANTU, 1971

Lithobius anisanus VERHOEFF, 1937

Synonyms: none

Lithobius austriacus VERHOEFF, 1937

Synonyms: Monotarsobius austriacus (VERHOEFF, 1937)


Synonyms: Lithobius lusitanus würmanus (VERHOEFF 1937); Lithobius lapidicola (LATZEL 1880)

Comment: The taxonomy of this species is

uncertain. It is only known from one record

from Selzthal in Styria and could possibly

be another species endemic to Austria.

Comment: Very closely related to

Lithobius aeruginosus.

43


Synonyms: Lithobius gridellii MANFREDI, 1955; Monotarsobius veronensis MATIC & DARABANTU,1971

Lithobius crassipes C. L. KOCH, 1862

Synonyms: Lithobius (Monotasobius) crassipes C. L. KOCH, 1862; Lithobius atrifrons SILVESTRI, 1896, Lithobius podokes ATTEMS, 1903

Lithobius curtipes C. L. KOCH, 1847

Synonyms: Lithobius carpathicus (MATIC 1958); Monotarsobius conformatus (CHAMBERLIN 1952)

Lithobius dentatus C. L. KOCH, 1844

Synonyms: Lithobius alpestris LATZEL, 1880


Synonyms: Lithobius armatus SSELIWANOFF, 1880; Lithobius dubius TÖMÖSVÁRY, 1880; Lithobius dubius TÖMÖSVÁRY, 1880

Comment: KOREN claims that the two

subspecies burzenlandicus and euxinicus

are recorded in Romania, while the

subspecies carinthicus is native to

Carinthia. Findings from Slovenia only

name the species with no subspecies

(KOS 1988). The findings of KOREN

generally match those made by MATIC in

1966. For the specimen found in Styria by

THALHAMER (1998) only the species is

given.

Comment: Originally described as L.

dubius by TÖMÖSVARY, in 1880, L.

erythrocephalus has been described by

many authors all throughout Europe. In

many places this species is distinguished

into two sympatric subspecies. The

subspecies L. erythrocephalus schuleri

(MATIC, 1966 schulleri) described by

VERHOEFF (1925, 1937) is much more

common in Austria, but the exact

taxonomy of the species and its

subspecies is in need of revision.

Lithobius forficatus (LINNÈ, 1758)

Synonyms: Lithobius leachii NEWPORT 1844; Lithobius vulgaris LEACH, 1817

Lithobius franzi ATTEMS, 1949

Synonyms: none


Synonyms: Lithobius pusillus (LATZEL 1880); Lithobius sulcatus (C. L. KOCH 1862)


Synonyms: Lithobius mutabilis transalpinus LATZEL, 1880


Synonyms: Lithobius alpinus C. L. KOCH, 1862; Lithobius rupivagus VERHOEFF, 1937; Lithobius walachicus VERHOEFF, 1901

Comment: The “common stone centipede”

is the most common species of centipedes

all throughout Europe and is omnipresent

in almost any habitat.

Comment: According to ATTEMS, L. franzi

is most closely related to L. walachius

(VERHOEFF, 1901) {= L. lucifugus C. L.

KOCH, 1862}. There is only one record of

this species and one type specimen, yet

its taxonomic status is unequivocally,

making it the only species endemic to

Styria and one of only two fully accepted

endemic centipedes of Austria.

45


Synonyms: Lithobius aulacopus LATZEL, 1880

Lithobius macrocentrus ATTEMS, 1949

Synonyms: none


Synonyms: Lithobius glabratus C.L. KOCH, 1847; Lithobius lusitanus crissolensis VERHOEFF, 1935


Synonyms: Lithobius daday TÖMÖSVÁRY, 1880; Lithobius communis C.L. KOCH, 1844; Lithobius maculatus MATIC & DARABANTU, 1971


Synonyms: Lithobius bicolor TÖMÖSVÁRY, 1879; Lithobius cinnamomueus L. KOCH, 1862

Comment: This endemic species has

been found and recorded multiple times in

Styria, Upper and Lower Austria as well as

Tyrol. The species is mostly alpine and

has been found over 3000 m above sea

level, yet has also been found in sub

montane areas just 500 m above sea

level.


Synonyms: Lithobius athesinus VERHOEFF, 1937

Lithobius peggauensis VERHOEFF, 1937

Synonyms: none


Synonyms: none


Synonyms: Lithobius marginatus FEDRIZZI, 1877; Lithobius fanzagoi FEDRIZZI, 1876

Lithobius pygmaeus LATZEL, 1880

Synonyms: Lithobius rucneri MATIC, 1966

Comment: Originally found and described

in Styrian Peggau, the species has also

been recorded in Slovakia and Bulgaria,

and its taxonomic status is highly

uncertain.

Comment: VERHOEFF noticed the variety of

certain traits of this speices and described

several subspecies: L. p. allemanicus from

Schwarzwald, L. p. insubricus from Wallis,

L. p. ponalensis from Gardasee and L. p.

annulipes from Hallstadt. A new

subspecies L. p. triangulatus nov. ssp. has

been described by KOREN and found at

the Saualpe.

47


Synonyms: Lithobius nigrifrons LATZEL & HAASE, 1880; Lithobius balcanicus MATIC, 1973


Synonyms: Lithobius dolomiticus ATTEMS, 1903; Lithobius oligoporus LATZEL, 1885


Synonyms: Lithobius leptotarsis MATIC, 1959; Lithobius molleri VERHOEFF, 1893; Lithobius punctulatus machadoi PRUNESCU, 1966

Lamyctes emarginatus (NEWPORT, 1844)

Synonyms: Lamyctes fulvicornis (MEINERT 1868); Lithobius gracilis (PORAT 1869)


Synonyms: Geophilus montanus (MEINERT,1870); Geophilus gorizensis (LATZEL,1880)

Comment: Is the only neobiotic species of

centipedes found in Austrias south (ESSL

& RABITSCH 2002). Its distribution is

primarily south-hemispheric but is often

recorded as Western-Palearctic (WÜRMLI

1972) (ZALESSKAJA & GOLOVATCH 1996)

and holarctic, and is indeed distributed

wide throughout the latter, and common in

tropical areas as well (ZALESSKAJA 1994).

It is a typical pioneering species in most

of Europe, preferring instable and

anthropogenic habitats of every kind. It

has been recorded in Styria by ATTEMS in

1895. There are no records in Carinthia

as of yet, but as the species kept

spreading in the last 20 years it should be

present in Carinthia as well.

Comment: According to VERHOEFF (1938)

the genus Clinopodes (ATTEMS 1929)

should be replaced by the genus

Geophilus, but Clinopodes remains as a

valid taxon. There are three subspecies

described: the “new” subspecies C. fl.

carinthicus LATZEL is synonymous with the

subspecies C. fl. flavidus ATTEMS. The

subspecies C. fl. styriacus ATTEMS 1895

and C. fl. polytrichus ATTEMS 1903 are

also native to Austria.


Synonyms: Geophilus brevicornis C.L. KOCH,1837; Himantarium caldarium MEINERT,1886; Geophilus ormanyensis ATTEMS,1903


Synonyms: Geophilus insculptus ATTEMS, 1895; Geophilus glacialis VERHOEFF, 1928; Geophilus glocknerensis VERHOEFF, 1940

Geophilus electricus (LINNÈ, 1758)

Synonyms: Geophilus sudeticus HAASE, 1880; Geophilus helveticus VERHOEFF, 1928


Synonyms: Necroploeophagus longicornis EASON 1960; Geophilus longicornis LEACH, 1815; G. pygmaeus styricus VERHOEFF, 1895


Synonyms: Geophilus noricus VERHOEFF, 1928; Geophilus pauropus ATTEMS, 1927; Geophilus insculptus ATTEMS, 1895


Clinopodes is being used by KOREN and

EASON but is invalid.

Comment: Old invalid names include

Geophilus linearis, Geophilus

glocknerensis, Geophilus proximus,

Geophilus glacialis, Geophilus anglicanus

and Geophilus alpinus.

Comment: Necroploephagus longicornis,

Geophilus longicornis, Geophilus

hortensis and Pachymerium tristanicum

are all old taxonomic names that are still

used but invalid.

Comment: Synonymous with the species

G. noricus (VERHOEFF 1928 and G.

pauropus (ATTEMS 1927), which have

been described as separate species by

KOREN and others.

49


Synonyms: Geophilus carnicus VERHOEFF, 1928; Geophilus cispadanus SILVESTRI, 1896; Geophilus larii VERHOEFF, 1934


Synonyms: Scolioplanes italicus VERHOEFF,1928; Linotaenia rosulans C.L. KOCH, 1847; Geophilus subtilis C.L. KOCH, 1838


Synonyms: Geophilus breviceps NEWPORT,1845; Scolioplanes mediterraneus VERHOEFF,1928


Synonyms: Scolioplanes transsilvanica (VERHOEFF, 1935)


Synonyms: Geophilus caucasicus ATTEMS,1903; Geophilus paradoxus TÖMÖSVÁRY,1880; Mecistocephalus punctilabium NEWPORT,184

Comment: According to ATTEMS 1929, G.

pygmaeus is an Illyric species native to

Slovenia, Croatia and Hungary. One

record from Carinthia and one from Styria

confirm the species is native to Austria. It

is also heavily implied that the species is

native to Upper Austria as well.


Synonyms: Schendyla (Echinoschendyla) zonalis BRÖLEMANN & RIBAUT, 1911


Synonyms: Dicellophilus nemorensis C. L. KOCH, 1837; Geophilus gracilis HARGER,1872


Synonyms: Schendyla montana ATTEMS, 1895; Brachyschendyla montana RIBAUT & BROLEMANN,1927


Synonyms: Geophilus apfelbecki VERHOEFF, 1898; Geophilus austriacus MEINERT, 1886; Mecistocephalus hungaricus TÖMÖSVÁRY,1880


Synonyms: Cryptops aenarienis VERHOEFF, 1943

Comment: The subgenus is

Echinoschendyla (BRÖLEMANN & RIBAUT),

but the old name Echinoschendyla zonalis

is invalid.


Dicellophilus is invalid.

Comment: The names Brachyschendyla/

Schendyla montana are not valid

anymore.

51


Synonyms: none

Scutigera coleoptrata (LINNÈ, 1758)

Synonyms: Scutigera pretzmanni WÜRMLI, 1973; Scutigera rubrovittata VERHOEFF, 1905

Comment: This species is the only repre-

senttative of the Scutigeromorpha in

Europe and very thermophile. It has

invaded almost every anthropomorphic

area thanks to carryover but has no high

abundances. It is extremely anthropo-

genic and common in houses and

manmade structures. There are only few

populations in the wild, which is why the

species is often listed as endangered.

KOREN (1992) and Zobodat© (2016) show

no records of Scutigera in Carinthia and

Styria, yet it is omnipresent. It is

discussed if the species should be

classified as a Neozoon, but experts claim

the species populated Austria from the

south by itself (ESSL &RABITSCH 2002).

Critical reflection

As many authors previously stated, most of the native centipede species are common. Within the 3rd, 4th

and 5th category, we will find 67,3 % of all species in Carinthia and 60,8 % in Styria. The fact that only 11-

19 % of all species native to Austrias south lack an efficient amount of data for statistical analysis, is once

again prove that the chilopod-fauna of the two counties Carinthia and Styria underwent far greater study

than the rest Austria as well as most of its neighboring countries.

The numbers of Carinthia and Styria match surprisingly well. While 9,6 % of Carinthian and 13,7 % of

Styrian species were declared “very rare” (Category 1), 11,5 % of Carinthian and 5,9 % of Styrian species

were declared as “rare” (Category 2), adding up to approximately 20 % of the species of both counties in

the bottom two categories. Same goes for categories (4) and (5), where Styria has more species in

category (5) and Carinthia more in category (4).

The terms “rare”, “omnipresent” and “common” are of course chosen and categorized within the

context of this study and the data available. As mentioned in the disclaimer, the fact that a species is

declared as “omnipresent”, does not directly indicate that the species can be found anywhere. A species

may still prefer a certain habitat or completely avoid another. A species declared “very rare” is not said

to be “endangered” in any way. It could be, but as previously mentioned, due to the lack of metadata on

populations and their growth no statements will be made about the species’ “vulnerability”. The species

Geophilus electricus (LINNÉ 1758) is a great example, being declared “rare” in Styria (Data = 8) and being

uncategorized in Carinthia (Data = 1). This species is well studied and nowhere near any endangerment

within Austria. But it can only be found in certain habitats underground, making it a very rare encounter.

Another curious case is Lithobius forficatus (LINNÉ 1758), the most common species of centipedes in most

central European countries. The species is listed in both counties as “omnipresent” yet has significantly

less findings than species like Cryptops parisi BRÖLEMANN 1921 and Strigamia acuminata (LEACH 1815),

two species that are definitely less “common” than forficatus, but might be encountered more often in

certain habitats. Another important factor, especially when comparing the amount of data between

Styria and Carinthia, is the fact that the south of Styria is not well studied compared to the rest of the

county and compared to Carinthia in its entirety. Another curious case, and possibly for the above

mentioned reason, is Clinopodes flavidus C. L. KOCH 1846. The species is quite common in Carinthia

(Category 4) and has been recorded many times but has only been found once in Styria in recent years in

the very south. The species is probably quite common in Styria (at least south of Graz), but could not

have been recorded more often, since the south has only undergone studies by centipede experts in

recent years. Findings from Carinthia imply that Clinopodes, although not completely absent, avoids

mountainous habitats as found in Styrias heavily studied north. Therefore, it is quite a task evaluating the

abundance of a species. Species like Harpolithobius anodus (LATZEL 1880) are vulnerable to some degree,

53

but the few relict populations we have are quite stable, so categorizing them as “very rare” might be

misleading if the list is interpreted the wrong way. On the other hand, a species like Lithobius pygmaeus

LATZEL 1880 (Category 0 in both Carinthia and Styria) is profiting from a lack of data in this case, since

only one finding more could put the species in Category 1 (very rare) which it is certainly not, but only a

very cryptic species with distribution mainly in mountainous areas.

Furthermore, evaluating a species with uncertain taxonomy is always a difficult task, especially if sources

disagree on the status of a species. Endemic and possibly endemic species are very important here. As of

now, there are two confirmed endemic species native to Styria. One of them, Lithobius franzi ATTEMS

1949 is endemic to Styria, the other, Lithobius macrocentrus ATTEMS 1949 has also been recorded in

Tyrol, Upper and Lower Austria and is endemic to Austria. The species Lithobius anisanus VERHOEFF 1937,

whose identity is completely uncertain, has only been found in the Styrian Selzthal and might also be

endemic to Styria. Same goes for the Carinthian Lithobius moellensis VERHOEFF 1940 so far, the only

species that might be endemic to Carinthia, whose identity is uncertain as well. When it comes to

neobiotic species, the situation in Austria is quite simple. The only species of centipedes declared

neobiotic is Lamyctes emarginatus (NEWPORT 1844), which has been recorded in Styria in recent years,

after spreading all over Austria from the east. The species has not been recorded in Carinthia yet but

might already be present and could be added to the list any day. The house centipede Scutigera

coleoptrata (LINNÉ 1758) has been declared neobiotic in many northern European countries by some

experts. In Austria (especially the south), the situation is more complicated. Though the species may be

highly anthropogenic and has not been found in the wild in neither Carinthia nor Styria, there are

populations of Scutigera in the northern parts of Austria as well as in the neighboring countries Slovenia,

Croatia and Italy. Experts claim Scutigera coleoptrata has colonized Carinthia and Styria from the south

on its own and has not established populations in the wild because of its anthropogenic preferences.

Summary and outlook

Overall, the lists give a good overview of the abundancy and ecology of the native species. Especially the

ecology of most of the species is well known and could assist in further evaluating a species’

endangerment, especially if its habitat is or might be subject to environmental changes. Each map also

gives a simple overview of the species’ geographical distribution and amount of data available. The

statistics show that most of the species are common and nowhere near any endangerment. For those

species declared “rare”, comments have been added to explain the “rarity”. Future friends of this taxon

should aim to improve upon this list and expand its categorization to species of other Austrian regions.

The difficulty of this task will of course heavily depend on how the taxonomic status of the native species

will change and whether or not the status of uncertain species will be cleared.

The easiest way to improve upon or add to the status quo is by collecting and determining individuals

and thereby gaining data, especially from those regions and habitats which have not undergone intense

(or any) studies so far (i.e. Styrias south). A specific search for cryptic Geophilomorpha or rare

Lihtobiomorpha would lead to an increase in data, thereby helping us better understand the biology,

distribution and possibly taxonomic history of the species, leading to an even more cohesive red list that

could tackle the difficult task of evaluating the vulnerability or even endangerment of at least a few

native species.

A work such as this is never fully finished. One can only aim to achieve greater goals. As the taxon of

centipedes is gaining more and more friends, and more people turning towards biodiversity and

ecological studies as a hobby in recent years, maybe as a collective we will be able to fully understand

the fascinating and often cryptic life of these versatile critters.

55

PART II.

DNA BARCODING OF CENTIPEDES

State of the art

For the German Barcode of Life initiative (GBOL), WESENER et al. (2015) had already experimented with

several primer pairs when trying to barcode Stenotaenia linearis (KOCH, 1835), a native species of

Geophilomorpha. As for over 200 Myriapoda species, the aim is to sequence parts of the mitochondrial

cytochrome c oxidase subunit I gene. For PCR and sequencing HCO/LCO primer pairs (FOLMER et al. 1994)

were utilized, which showed a low success rate (<50%). The degenerate primer pair HCOJJ/LCOJJ (ASTRIN

& STÜBEN 2008) was used for further sequencing attempts, resulting in a much higher success rate

(>75%). In 2011 SPELDA et al. pioneered research in this field in Germany and sequenced many Bavarian

species of myriapods for the first time.

According to the official GBOL website 37 species of Chilopoda have been barcoded by the project since

its initiation (Dec. 2019), with 658 individuals registered in the database, some not determined further

than at family level (and some missing with actual barcode data). 658 individual centipedes have been

barcoded within the GBOL project including the following species: Lithobius forficatus (LINNÉ 1758),

Lithobius erythrocephalus C. L. KOCH 1847, Lithobius piceus C. L. KOCH 1862, Lithobius mutabilis C. L. KOCH

1862, Lithobius tenebrosus MEINERT 1872, Lithobius agilis C. L. KOCH 1874, Lithobius melanops NEWPORT

1845, Lithobius austriacus (VERHOEFF 1937), Lithobius malicentus C. L. KOCH 1862, Lithobius tricuspis

MEINERT 1872, Lithobius dentatus C. L. KOCH 1844, Lithobius muticus C. L. KOCH 1847, Lithobius borealis

MEINERT 1868, Lithobius microps MEINERT 1868, Lithobius crassipes C. L. KOCH 1862, Lithobius pelidnus

HAASE 1880, Lithobius curtipes C. L. KOCH 1847, Lithobius calcaratus C. L. KOCH 1844, Lithobius nodulipes

LATZEL 1880, Lithobius aeruginosus C. L. KOCH 1862, Eupolybothrus tridentinus (FANZAGO 1874), Cryptops

hortensis (DONOVAN 1810), Cryptops anomalans NEWPORT 1844, Cryptops parisi BRÖLEMANN 1921,

Geophilus truncorum BERGSØE & MEINERT 1866, Geophilus alpinus MEINERT 1869, Geophilus carpophagus

LEACH 1816, Geophilus flavus (DE GEER 1778), Geophilus electricus (LINNÉ 1758), Stenotaenia linearis (C. L.

KOCH 1835), Pachymerium ferrugineum (C. L. KOCH 1835), Schendyla nemorensis (C. L. KOCH 1837),

Stigmatogaster subterranean (SHAW 1974), Lamyctes emarginatus (NEWPORT 1844), Strigamia acuminata

(LEACH 1815), Strigamia crassipes (C. L. KOCH 1835), Henia vesuviana (NEWPORT 1845).

For the Austrian Barcode of Life (ABOL) project Dr. Nesrine Akkaris work group already sequenced

several species of centipedes, mainly of the genus Lithobius including the species’ Lithobius nodulipes

LATZEL 1880, Lithobius pelidnus HAASE 1880, and Lithobius pygmaeus LATZEL 1880 from Carinthia, with the

four markers COI, 16S, 18S and 28S6. An extensive study by OEYEN et al. (2014) unveiled the evolutionary

history of the Austrian Scolopendra cingulata LATREILLE 1829 population. The large gap between the intra-

and interspecific distances not only confirmed the Austrian and Hungarian populations to be relict

populations of a much wider distribution during the last post-glacial climatic optimum (which later

became isolated by the spreading of forests), but also shows that the Austrian population has a

completely unique haplotype within the species, despite its low genetic variation. A similar study by

WESENER et al. (2016), focusing on barcoding Centipedes from Austria and Germany, revealed large

intraspecific distances in several Cryptops species, with German specimens formally assigned to Cryptops

parisi BRÖLEMANN 1921 divided into three clades differing by 8.4–11.3% from one another. This study also

revealed the existence of several ghost lineages and revealed new species for Germany such as Cryptops

umbricus VERHOEFF 1931.

For this thesis 32 different species have been barcoded with the COI maker from different Austrian

areas, for 26 of which at least one barcode was successfully generated. Scolopendra cingulata LATRILLE

1829 and Geophilus electricus (LINNÉ 1758) as well as two unidentified Lithobius specimens were

collected in the state of Burgenland. From the same area in Burgenland as well as from the FH Graz

Campus the species Scutigera coleoptrata (LINNÉ 1758) has been barcoded, Pachymerium ferrugineum (C.

L. KOCH 1835), Lithobius erythrocephalus C. L. KOCH 1847, Lithobius aeruginosus C. L. KOCH 1862, Lithobius

latro MEINERT 1872,Lithobius pelidnus HAASE 1880, and Lithobius agilis C. L. KOCH 1874 from Lower

Austria, Lithobius lucifugus C. L. KOCH 1862 from Lower Austria and Carinthia, Eupolybothrus grossipes (C.

L. KOCH 1847), Lithobius macilentus C. L. KOCH 1862, Lithobius mutabilis C. L. KOCH 1862 and Schendyla

tyrolensis ATTEMS 1895 from Carinthia, Eupolybothrus tridentinus (FANZAGO 1874), Lithobius muticus C. L.

KOCH 1847, Schendyla carniolensis VERHOEFF 1902 and Cryptops hortensis (DONOVAN 1810) from Styria,

Henia illyrica (MEINERT 1870) and Henia vesuviana (NEWPORT 1845) from Vienna, Lithobius tricuspis

MEINERT 1872 from Carinthia, Clinopodes flavidus C. L. KOCH 1847 from Carinthia and Vienna, Strigamia

acuminata (LEACH 1815) from Styria, Vorarlberg and Upper Austria, Strigamia transsilvanica (VERHOEFF

1928) from Styria, Vorarlberg and Upper Austria, and from several counties the more widespread species

Lithobius dentatus C. L. KOCH 1844, Lithobius forficatus (LINNÉ 1758), Lithobius piceus C. L. KOCH 1862,

Lithobius tenebrosus MEINERT 1872, Lithobius validus MEINERT 1872, Dicellophilus carniolensis (C. L. KOCH

1847) and Cryptops parisi BRÖLEMANN 1921.

6 Data yet unpublished

57

Austrian centipede species that already underwent barcoding include Lithobius forficatus (LINNÉ 1758),

Lithobius dentatus C. L. KOCH 1844, Lithobius schuleri VERHOEFF 1925, Lithobius agilis C. L. KOCH 1874,

from Upper Austria, Lithobius glacialis (ATTEMS 1909), Strigamia acuminata (LEACH 1815) and Cryptops

parisi BRÖLEMANN 1921 from Upper Austria and Salzburg, Lithobius piceus C. L. KOCH 1862, Lithobius

tenebrosus MEINERT 1872, Lithobius aeruginosus C. L. KOCH 1862, Lithobius austriacus (VERHOEFF 1937),

Geophilus alpinus MEINERT 1870, Geophilus flavus (DE GEER 1778) and Schendyla tyrolensis (MEINERT 1870)

from Salzburg and Scolopendra cingulata LATRILLE 1829 from Burgenland, as well as 25 other species only

identified as part of the Lithobiomorpha and an additional six identified as Geophilomorpha order from

Salzburg, Upper Austria, Carinthia and Tyrol.7 Barcodes of species from Styria, Lower Austria and Vienna

(which are included in this thesis) are completely absent. Of the barcode data available on BOLD© 48

specimens are from the institution Zoologische Staatssammlung in Munich, the data of the remaining 10

specimens is retrieved from GenBank©.

7 All with COI markers; source: boldsystems.org (March 2020)

Methodology

The centipedes barcoded for this thesis were both collected by me and other researchers. Many

specimens were provided by Nikolaus Szucsich of the NHM Vienna. A complete list of all individuals,

collectors and locations can be found in the appendix.

Literature for species determination of centipedes is sparse, especially those with depictions of the

morphological structures (most only show representative drawings of the last pair of legs). Fortunately

KOREN wrote some of the best determination keys for the Geophilomorpha & Scolopendromorpha (1986)

and Lithobiomorpha (1992) for Carinthia and Eastern Tyrol, based upon and including drawings from the

works of EASON (1964). These determination keys also apply to most of the species found in Styria, since

there are only a handful species present in Styria that are absent in Carinthia (see Chapter I). For a

couple of other species, determination was done using internet research, individual papers or

descriptions and depictions of the species found in online centipede forums.

The critters were stored in 100% Ethanol in small test tubes. Some individuals have previously been

stored in 70% Ethanol but were then relocated to pure Ethanol. Individual names were attached to the

samples consisting of a code which contained the species name, sample number and location of

collection of the individual. For example, the individual number 51 is the species Lithobius forficatus

(LINNÉ 1758) and was the 4th individual collected at the location “Johnsbach”. The resulting code for the

species is 51_FOR_JB4.

Mitochondrial DNA has been extracted in the laboratory facilities of the Department of Biology.

Extraction of the material (mostly stored in 100% EtOH) was achieved by cutting off small pieces (i.e. one

leg) of the centipedes, drying the piece quickly thereby draining it from the alcohol and then placing it in

10% Chelex® solution, as was done by RICHLEN & BARBER (2005). The vortexed and centrifuged material

then underwent the extraction process for 20 minutes at 95°C.

For PCR the primer pair LCO1490‐JJ2/HCO2198‐JJ2 (ASTRIN & STÜBEN 2008) has been used, which has

shown great success in barcoding centipedes for the German GBOL project in recent years. Three

different types of Taq-polymerase had been used in the process: PhusionTM, BioThermTM and

SupraThermTM, all to great success with PhusionTM generally being the most effective and BioThermTM

the least. After trying out different annealing temperatures between 48°C and 51°C during the PCR

process results varied as well. DNA amplification followed the protocol of DUFTNER et. al 2005. The PCR

cocktail (9 µl) consisted of a Taq-polymerase (0,2 µl for PhusionTM/ 0,1 µl for Bio/SupraThermTM), the

primer pair (1 µl/ 0,5µl), dNTP’s (0,5 µl/ 0,35 µl), water (5,3 µl/ 7,5 µl) and a corresponding buffer (2 µl/ 1

µl), mixed with 2 µl of the sample. PCR was performed at a maximum temperature of 95°C and either

60°C or 72°C at the end of each cycle. The number of cycles varied as well, as 35 is the maximum number

59

for PhusionTM and 45 cycles the regular amount for primers such as BioThermTM and SupraThermTM

(KOBLMÜLLER et. al 2011).

After electrophoresis, samples with clearly visible bands underwent cleanup via ExoSAPTM, a reagent

which hydrolyzes excess primer fragments and nucleotides, followed by the sequencing reaction. For the

three-hour sequencing reaction at 94°C initial temperature and annealing temperature of 50°C, a 6 µl

cocktail of H2O (3,95 µl), sequencing buffer (1,6 µl), BigDye® terminator (0,2 µl) and the primer (0,25 µ)

was mixed with 2 µl of the sample (KOBLMÜLLER et. al 2011). After adding distilled water to the samples

and pipetting them onto SephadexTM pillars inside Eppendorf test-tubes, the samples were centrifuged at

9000rpm and thereby purified. What followed was the sequencing process using a 16-capillary-

sequencer 3130x and a 3500xl 24-capillary-sequencer by Applied Biosystems®. The resulting sequences

were (if necessary) corrected, inserted into a FASTA file and aligned using the MEGA7© program. After

shortening all the sequences to “upload length” the sequences were “blasted” using the BOLD©

(www.boldsystems.org) database, and matching sequences were downloaded from GenBank©

(https://www.ncbi.nlm.nih.gov/genbank) to be inserted into the FASTA file for comparison and further

analysis.

From there a Neighbor-Joining tree could be constructed using MEGA©, a agglomerative bottom-up

clustering method that requires knowledge of the distances between each pair of taxa and is commonly

used when working with protein-coding DNA data. A p-distance model has been chosen for substitution

with uniform rates and a 1000 bootstrap method for statistical support. In order to further estimate the

evolutionary divergences between the sequences/species, a matrix output was created displaying

between group mean distances. Maximum intraspecific and minimum interspecific differences were

evaluated manually using Microsoft Excel©. The same file was then used to find the nearest neighbor of

those species which were to further evaluate, i.e. those where data of three or more specimens was

available. For those species a distance sheet was created.

http://www.boldsystems.org/

https://www.ncbi.nlm.nih.gov/genbank

https://en.wikipedia.org/wiki/Cluster_analysis

Results and discussion

Graphic 2: Complete neighbor-joining tree computed using MEGA7.

61

Graphic 3: Circle NJ tree of the Geophilomorpha computed using MEGA7.

Graphic 4: Circle NJ tree of the Lithobiomorpha computed using MEGA7.

Graphic 5: NJ tree of the Scolopendromorpha and Scutigeromorpha computed using MEGA7.

Table 2: Interspecific distances and distances to nearest neighbor (DNN) evaluated for every species with n>=3.

Species N Imax Nearest Neighbour DNN

Scutigeromorpha

Scutigeridae

Scutigera coleptrata 3 4,3 Cryptops hortensis 22,2

Scolopendromorpha

Cryptopidae

Cryptops parisi 4 9,6 Cryptops anomalans 18,4

Lithobiomorpha

Lithobiidae

Lithobius erythrocephalus 3 15,4 Lithobius piceus 12,9

Lithobius forficatus 5 14,5 Lithobius validus 16,6

Lithobius tenebrosus 3 9,3 Lithobius piceus 14,9

Lithobius validus 5 13,7 Lithobius piceus 15,6

Geophilomorpha

Geophilidae

Strigamia acuminata 9 15,7 Strigamia transsilvanica 15,4

Henia illyrica 4 11,6 Henia vesuviana 8

Henia vesuviana 4 3,9 Henia illyrica 8

63

The complete NJ-tree (Graphic 2, p. 60) shows a clear distinction between the orders Lithobiomorpha

and Geophilomorpha. The Scutigeromorpha are falsely potted with the Scolopendromorpha, rather than

being an outside group. COI barcoding data is however unreliable for evaluating and supporting larger

phylogenetic splits due to the high mutation rate of the gene, and should only be used to distinguish

closely related species. With the exception of Pachymerium ferrugineum (C. L. KOCH 1835) all members of

the Geophilomorpha order can be found potted at the bottom of the tree. The odd positioning of

ferrugineum as an outside group of the Geophilomorpha is even more evident in the Geophilomorpha

NJ-tree (Graphic 3, p. 61), but can (again) be explained away with the limitations of the barcoding data

when it comes to supporting deeper phylogenetic splits. While Pachymerium may not be closely related

to other Geophilomorpha genera such as Strigamia it is usually found closer to genera such as Geophilus.

The unknown Lithobius specimen #1 seems to be related to Lithobius forficatus (LINNÉ 1758) but doesn’t

appear to be Lithobius piceus C. L. KOCH 1862, which it shared many morphological traits with. Judging

from the tree alone, the 2nd unidentified Lithobius specimen is most probably Lithobius macilentus C. L.

KOCH 1862, a morphological analysis however strongly suggests otherwise.

The notably high distances displayed in Table 2 (p. 62) suggest high levels of cryptic diversity. Studies on

cryptic diversity in centipedes are not sparse, but usually restricted to a confined area such as islands or

archipelagos. Cryptic diversity of central European centipedes has mostly been studies by WESENER,

VOIGTLÄNDER and their associates. They all have come to conclusion that cryptic diversity in European

centipedes is not uncommon, especially in older and omnipresent species like Cryptops parisi BRÖLEMANN

1921 (WESENER et. Al 2016). The highest intraspecific distance in Lithobius erythrocephalus C. L. KOCH

1847 strangely enough appears to be lower than the distance to its nearest neighbor Lithobius piceus C.

L. KOCH 1862. The two specimens closest to each other were retrieved from GenBank, however the

distance to the other piceus specimens are all lower than the imax for erythrocephalus as well. Lithobius

erythrocephalus C. L. KOCH 1847 is a species with several subspecies in Europe, two of which are native to

Austria. Even KOREN (1992) noted that the status of erythrocepahlus and all its subspecies should be

evaluated in order to track the species’ history. Lithobius piceus C. L. KOCH 1862 is an older species and

omnipresent all throughout Europe. It is therefore probably very diverse as well. We face the same

situation with Henia illyrica (MEINERT 1870) and Strigamia acuminata (LEACH 1815). The imax of the latter

however almost matches the distance to its nearest neighbor Strigamia transsilvanica (VERHOEFF 1928).

Both species show high levels of diversity with distances ranging from 0.2 to 15.7. The species’ of the

genus Henia are known to be very closely related to each other. The high intraspecific distance of Henia

illyrica (MEINERT 1870) can be explained by the inclusion of one specimen retrieved from GenBank©

which was collected in Germany. The large geographic distance is reflected in the genetic distance,

further underlining the diversity of these critters. Greater interpretation of the data and results is not

easy due to the low number of specimens per species.

Summary and outlook

The majority of the laboratory analysis has been carried out between February and June of 2020 and

therefore has been interrupted for weeks due to Corona-Virus safety measures, when university facilities

remained closed to all non-essential staffers. DNA barcoding has been largely unsuccessful ever after the

end of the regulations in late May. The specimens have been extracted several times, primers and

polymerases were changed and experimented with in order to achieve satisfying results, yet none of the

trials proved successful. The ultimate decision was to work with what we had. Unfortunately, the lack of

a substantial amount of usable barcodes per species and location left us unable to upload any barcodes

to the ABOL data bank or GenBank©, as was previously one of the goals of this thesis. However outside

of this thesis laboratory work may continue in order to create publishable barcodes.

Today several authors note that species identification using morphological traits has several limitations

such as misidentification due to plasticity of a trait or the existence of cryptic taxa. A high level of

expertise is often required to correctly identify species with the required certainty, and even the best

morphological keys are often only of use in one particular life stage or gender if based on genitalia. The

DNA barcoding approach might currently represent the best solution for species identification, even if

barcoding itself represents some limitations (VALENTINI et al. 2008, SALVI et. al 2020). Identifying species

through DNA barcoding is also helpful for understanding interspecies interactions as researchers such as

PFENNINGER et al. (2007) and TEDERSOO et al. (2007) point out. The number of studies involving DNA

barcoding has risen sharply ever since 2017, with the rise of newer and more effective methods which

(by then) have become broader available (DE SALLE & GOLDSTEIN 2019). DNA barcoding has already led to

many important phylogenetic discoveries, as barcoding of Austrian Scolopendra cingulata LATREILLE 1892

populations by OEYEN et al. (2014) revealed the population to be an important biogeographical relic in a

possible microrefugium.

DNA barcoding is mainly limited by the system it is using, which is single locus based, even if several

regions from the organelles DNA are sequenced. It is known and has been well discussed that identical

mitochondrial DNA sequences can be present in different related species, either due to introgression or

incomplete lineage sorting since the time of speciation (BALLARD & WHITLOCK 2004). The inclusion of

mtDNA evidence in biogeographic analysis often reveals unexpected diversity or discordance with

morphology, prompting a re-evaluation of morphological and ecological characteristics and even

taxonomic revision (MORITZ & CICERO 2004). MORITZ & CICERO (2004) point out that mtDNA divergence

should not be a primary criterion for recognizing species boundaries and further claim: “[…] focusing on

mtDNA divergence as a primary criterion for recognizing species […] may lead scientist to overlook new

or rapidly diverged species, such as might arise through divergent selection or polyploidy […]”. There are

several other problems such as the accidental amplification of nuclear copies of mitochondrial DNA

65

fragments and the use of a divergence threshold for intra- & interspecies variation distinction (ZHANG &

HEWITT 1996, HEBERT et al. 2003). According to NIELSEN & MATZ (2006) several cases of species

misidentification are caused by incomplete databases, where the number of analyzed individuals per

species is often far too low and does not allow for a precise estimation of intra- and interspecific

variation. HEBERT et al. (2003) mentioned another limitation of DNA barcoding in in the length of the

sequences used, which is usually > 500 bp and thereby prevents the amplification of degraded DNA,

which many DNA barcoding applications can only be based on, especially environmental samples where

the target DNA is often from dead animals or plants (DEAGLE et al. 2006, VALENTINI et al. 2008). The rise of

NGS brought fourth even newer, more revolutionary barcoding methods such as “mini-barcoding”,

methods that are used to this very day. VALENTINI et al. in 2008 proposed NGS methods such as

pyrosequencing could counteract many of the upper mentioned problems; however pyrosequencing did

not see the rise the authors expected and was quickly replaced by cheaper NGS methods.

With many methods failing to provide cohesive results a lot of doubt has been cast on the DNA

barcoding method in the late 2000’s, yet ecologists increasingly turned to barcoding methods as a means

of species determination, because in many circumstances it represents an easy and rather quick way to

identify a species. HULLEY et. al (2018) found another use of DNA barcoding in their research on larval fish

in the great lakes. They deemed DNA barcoding: “an effective alternative to morphological identification

[…] of larval and embryonic fish, but comparisons of the two approaches with species from the Great

Lakes are limited. It may be particularly important to examine this issue in the Great Lakes because a

relatively young group of post-glacial fish species are present which may be difficult to resolve using

morphology or genetics.” Consequently both critics and champions of the barcoding method published a

wide variety of papers on the subject in the last decade. General interest in barcoding, as well as careful

and critical analysis, led to the rise of more effective methods and even approaches specific to one group

of organisms or even a single genus. Still, the accuracy of DNA barcoding tools depends on the existence

of a comprehensive archived library of sequences reliably determined at species level by expert

taxonomists, as SALVI et. al (2020) point out. Pre-barcoding morphological analysis is especially important

when working with a new or small group of animals, where new subspecies or variates are expected to

still be discovered. It is most important to always have a DNA barcode of the sample as basic data, since

the sequences are not prone to subjectivity and can be reanalyzed in the future in accordance with

improvements in taxonomic knowledge, as VALENTINI et al. (2008) point out. SALVI et. al (2020) further

discussed the limitations of barcoding and the issue of intrinsic and extrinsic errors when they explain:

“While intrinsic errors to be solved require an integrative taxonomic and evolutionary study approach,

which goes beyond the idea of barcoding as identification tool, extrinsic errors are due to human

mistakes and can be corrected much more easily. Thus, in a reference dataset free of extrinsic errors it

would be easy to spot species identification inconsistencies that require further taxonomic research, [...]

https://www.sciencedirect.com/topics/earth-and-planetary-sciences/postglacial

the importance of which [...] cannot be overstated”. While DE SALLE & GOLDSTEIN (2019) seem to agree

with most researchers defending the DNA barcoding method as a valuable taxonomic tool, they very

critically discuss distance measures and NJ trees derived from barcoding data as a means of species

determination. They claim: “it is important to separate the statement that NJ analyses “work” to identify

species from the supposition that they allow us to infer anything about species in the abstract” and

summarize a long discussion with: “[...] the fact that a very high proportion of diagnosable species are

captured by NJ analyses is encouraging, but not sufficient.” Thereby they reject the premise that DNA

barcoding serves to repair some inherent flaw in the practice of systematic, yet actively embrace DNA

barcoding as a near-universal advance for taxonomic research.

Generally speaking, DNA barcoding works well in most species, especially when the amount of data is

large and covers the total variability of the species’ area of distribution. Problems may still occur with

either very young species that did not accumulate or sort out enough mutations to distinguish them by

their barcodes alone or old species with large population sizes where divergent haplotypes may survive.

Hybridization and introgression events may still lead to “wrong barcodes”, but refined methods aim to

avoid those problems (ZIMMERMANN et. al 2013). As is evident, the method is an effective and powerful

one, and should therefore be used with great care and thought put into it. A critical analysis of one’s own

methods is fundamental in evolutionary biology.

As far as centipedes are concerned, the DNA barcoding approach alone will not resolve their taxonomy.

The high levels of cryptic diversity, numerous subspecies and high variability of important traits (such as

gonopods shape, number of segments etc.) make a morphological (pre)analysis essential. DNA barcoding

should however be applied whenever possible, since it is an important tool that may aid in taxonomic

determination. Another acceptable approach for example would be to simply generate barcodes with

very little pre-analysis in order to search for signs of cryptic diversity. This approach might also reveal

other details of greater importance, such as the revelations of the research by OEYEN et al. (2014) on

native Scolopendra populations, which would have remained completely unobserved without the usage

of the barcoding method.

The results of this very thesis have shown what issues may occur. The barcode of one individual may

match the barcode of a second individual, yet morphologically they can be completely different. The

problems we (possibly) face when applying or relying on the barcoding method are various and have

been discussed extensively above. Problems should always be critically examined individually. Many

taxonomists face the problem of numerous subspecies within their orders of expertise. Discussions on

the actual existence of subspecies as biological units will continue and not be resolved any time soon.

Whatever these subspecies may represent, they are proof of high levels of variety within certain species’.

As we have shown, DNA barcoding may assist in evaluating the gaps between these varieties. Knowledge

67

about the species’ biology, ecology, morphology etc. is fundamental, because it aids us in interpreting

the barcoding results and helps resolving possible issues and mismatches.

Lastly, DNA barcoding will continue to evolve and improve. We cannot predict what new methods will

arise in the future, but we shall aim to improve upon what is established. Solving the taxonomy of all

beings sharing this world with us is a massive task, but (as is evident) many scientists take on this task

with more and more joining every day. With combined efforts and many friends of different organisms

we can aim to achieve greater knowledge about the history of all kinds of organisms with the help of

present and future DNA barcoding methods.

APPENDIX

List of individual centipede specimens included in the tree

Specimen# Species Collection Site Coordinates Date o. Coll. Collector Determinatior

6_COL_TR3 Scutigera coleoptrata Thenauriegel (BL) 47.941453, 16.718001 08.06.2018 Svetnik Svetnik

48_COL_FH1 Scutigera coleoptrata FH Joanneum Graz (Stmk) 47.069168, 15.410829 04.07.2019 Lienhard Svetnik

43_VAL_E12 Lithobius validus Leechwald Graz (Stmk) 47.084431, 15.462240 01.04.2019 Svetnik Svetnik

66_VAL_X1 Lithobius validus Tainach (Ktn) 46.391780, 14330349 14.04.2017 Svetnik Svetnik

67_VAL_X2 Lithobius validus Gut Landskron (Ktn) 46.645287, 13.895456 27.07.2019 Svetnik Svetnik

68_VAL_BL6 Lithobius validus Burg Landskron (Ktn) 46.641788, 13.898047 07.08.2018 Svetnik Svetnik

36_FOR_E1 Lithobius forficatus Leechwald Graz (Stmk) 47.084431, 15.462240 01.04.2019 Svetnik Svetnik

45_FOR_NO9 Lithobius forficatus Schönau (NÖ) 48.135400, 16.613800 29.04.2018 Wagner, Spiß Svetnik

51_FOR_JB4 Lithobius forficatus Johnsbach (Stmk) 47.321630, 14.372630 19.06.2018 Svetnik Svetnik

56_FOR_NO6 Lithobius forficatus Schönau (NÖ) 48.135400, 16.613800 29.04.2018 Wagner, Spiß Svetnik

53_PIC_JB6 Lithobius piceus Johnsbach (Stmk) 47.321630, 14.372630 19.06.2018 Svetnik Svetnik

87_PIC_1960 Lithobius piceus* St. Pölten (NÖ) 47.995619, 15.456727 unknown Bartel, Böhm Svetnik

107_DEN_2330 Lithobius dentatus Johnsbach (Stmk) 47.569639, 14.583405 unknown Macek, Bartel Svetnik

27_TEN_E6 Lithobius tenebrosus Leechwald Graz (Stmk) 47.086294, 15.462648 01.04.2019 Svetnik Svetnik

58_TEN_WH17 Lithobius tenebrosus Waldhof Villach (Ktn) 46.636078, 13.896829 14.08.2019 Svetnik Svetnik

15_BOR_WF10 Lithobius borealis Waldhof Villach (Ktn) 46.635511, 13.896883 04.08.2018 Svetnik Svetnik

70_MAL_WF11 Lithobius macilentus* Waldhof Villach (Ktn) 46.635511, 13.896883 04.08.2018 Svetnik Svetnik

112_IND_2366 Lithobius indet. Gsuchmauer (Stmk) 47.547383, 14.656950 unknown Group NHM Svetnik

23_ERY_NO8 Lithobius erythrocephalus Schönau (NÖ) 48.134100, 16.606900 29.04.2018 Wagner, Spiß Svetnik

91_ERY_1955 Lithobius erythrocephalus St. Pölten (NÖ) 47.995619, 15.456727 unknown Bartel, Böhm Svetnik

83_IND_2193 Lithobius indet. Ruine Neuburg (Vbg) 47.423219, 9.622580 unknown Macek Svetnik

86_GLA_1980 Lithobius glacialis St. Pölten (NÖ) 47.914555, 15.372111 unknown Macek, Bartel Svetnik

26_HOR_E9 Cryptops hortensis Leechwald Graz (Stmk) 47.084431, 15.462240 01.04.2019 Svetnik Svetnik

46_ANO_NO10 Cryptops anomalans Schönau (NÖ) 48.135400, 16.613800 29.04.2018 Spiß Svetnik

42_PAR_E11 Cryptops parisi Leechwald Graz (Stmk) 47.084431, 15.462240 01.04.2019 Svetnik Svetnik

65_PAR_AB1 Cryptops parisi Gut Landskron (Ktn) 46.644767, 13.899715 27.07.2019 Svetnik Svetnik

88_PAR_1956 Cryptops parisi St. Pölten (NÖ) 47.995619, 15.456727 unknown Bartel, Böhm Svetnik

96_ELE_2235 Geophilus electricus Ebner Klamm (Stmk) 47.530836, 14.530836 unknown Macek, Bartel Svetnik

25_ACU_E14 Strigamia acuminata Leechwald Graz (Stmk) 47.086294, 15.462648 04.05.2018 Svetnik Svetnik

76_ACU_2044 Strigamia acuminata Tschütsch (Vbg) 47.315355, 9.646802 unknown Macek Svetnik

77_ACU_2050 Strigamia acuminata Krummberg (Vbg) 47.334980, 9.606916 unknown Macek Svetnik

84_ACU_2177 Strigamia acuminata Kloster Maria Stern (Vbg) 47.570386, 9.779252 unknown Macek Svetnik

85_ACU_2078 Strigamia acuminata Kloster Maria Stern (Vbg) 47.570386, 9.779252 unknown Macek Svetnik

97_ACU_2251 Strigamia acuminata Ebner Klamm (Stmk) 47.530836, 14.530836 unknown Macek, Bartel Svetnik

98_ACU_2267 Strigamia acuminata Ödelsteinhöhle (Stmk) 47.519166, 14.615277 unknown Macek, Bartel Svetnik

102_ACU_2274 Strigamia acuminata Ödelsteinhöhle (Stmk) 47.519166, 14.615278 unknown Macek, Bartel Svetnik

111_TRA_2358 Strigamia transsilvanica Landweg Liezen (Stmk) 47.609808, 14.740686 unknown Macek, Bartel Svetnik

71_ILL_1227 Henia illyrica Baumgartner Höhe (W) 48.210816, 16.277955 unknown Bartel, Szucsich Szucsich

74_ILL_1528 Henia illyrica Döbling (W) 48.275700, 16.351986 unknown Szucsich Szucsich

95_ILL_1599 Henia illyrica Prater (W) 48.191666, 16.445000 unknown Szucsich Szucsich

72_VES_1321 Henia vesuviana Wienflussufer (W) 48.209833, 16.212641 unknown Bartel, Szucsich Szucsich

73_VES_1442 Henia vesuviana Schlosspark Schönbrunn (W) 48.183330, 16.300000 unknown Szucsich Szucsich

81_VES_1443 Henia vesuviana Schlosspark Schönbrunn (W) 48.183330, 16.300001 unknown Szucsich Szucsich

94_NEM_2196 Schendyla nemorensis Ruine Neuburg (Vbg) 47.423219, 9.622580 unknown Macek Svetnik

78_TYR_2214 Schendyla tyrolensis Wolfsberg Graben (Ktn) 46.898333, 14.848888 unknown Walzl Svetnik

82_FLA_1530 Clinopodes flavidus Döbling (W) 48.275700, 16.351986 unknown Szucsich Szucsich

37_FER_NO1 Pachymerium ferrugineum Hafen Lobau (NÖ) 48.154200, 16.487600 30.04.2018 Wagner, Spiß Svetnik

41_CAR_E10 Dicellophilus carniolensis Leechwald Graz (Stmk) 47.084431, 15.462240 04.05.2018 Svetnik Svetnik

*reverese Taxonomy

69

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