Giant pill-millipede keeping :

courtesy to : www.sanbi.org/creature/giant-pill-millipede

Derivation of scientific name :

The name millipede comes from the Latin word mille meaning thousand andpedis meaning foot. Even though these animals are called millipedes, they don’t have a thousand legs; most of the species have between 36 and 400 legs with the exception of the rare species called Illacme plenipes, which was recorded to have 750 legs. The order name Sphaerotheriida comes from the Latin word sphaerium, meaning ball, which reflects the ability of pill-millipedes to roll up into a ball.

The giant pill-millipede belongs to a unique group of millipedes that differ in appearance from the usual long, worm-like and numerous-legged millipedes that most people are familiar with. Instead, pill-millipedes are characterised by an unusual round and stout body profile with few inconspicuous legs. The body is covered by a strong exoskeleton comprised of curved shields (tergites) that are used as a passive defensive mechanism when threatened. Pill-millipedes are able to roll themselves into a tight ball with the vulnerable soft underbelly tucked inside and protected by the outer, tough exoskeleton. In order to attract females for breeding, male pill-millipedes produce sounds and vibrations through an act known as stridulation. The giant pill-millipede is ground dwelling and occurs in moist habitats under leaf litter on the forest floor and under trees. It is widely distributed in South Africa with records from the higher-altitude, forested parts of Limpopo, KwaZulu-Natal and the Eastern Cape. Currently, pill-millipedes in South Africa are not formally protected and they may be vulnerable to habitat loss and degradation, as well as overexploitation for use in traditional medicine.

Description:

The giant pill-millipede is the largest species of pill-millipede that occurs in South Africa. Its total length when unrolled and walking can be as long as 5 cm, and when rolled up, the giant pill-millipede can be as large as a ping-pong ball. The body is short, stout and composed of 11–13 segments and each segment has two pairs of legs. It has one pair of antennae and a pair of kidney-shaped eyes. The body segments are specifically designed to be able to completely encase the millipede when it rolls itself up into tight ball. The head and first segment are very small and can be easily tucked in under the very large second segment that has a raised rim that closely fits and engages the last segment when rolled up. The last segment is large and has a distinctive appearance of a half bell. The giant pill-millipede is chestnut brown with black posterior edges to each segment, giving it a brown and black banded appearance.

Getting around:

Giant pill-millipedes, like other members of the genus, have about 42 legs and this, together with their wide body and heavy cuticle, cause them to move very slowly. Giant pill-millipedes are also able to burrow into the soil and leaf litter.

Communication :

Sphaerotherium species are unusual in their ability to produce sound and vibrations by rubbing together certain body parts, an act known as stridulating. This is normally done by males during mating. In males the last two pairs of legs on the last segment are modified and are called telopods (clamp-like structures). These telopods have an area with a series of sclerotised ridges that are rubbed against knobs on the underside of the anal shield. Each species has a different number and width of ridges on the telopods, and so the stridulating sound and vibrations produced are unique to each species. The female can therefore detect suitable mates based on the vibrating patterns. The specific frequency of the stridulation seems to stop the female from rolling up and allow the male to mate with her. Since pill-millipedes have no hearing organs, they must be picking up vibrations rather than hearing the sound.

Distribution :

The giant pill-millipede is widely distributed in South Africa with records from Limpopo, Mpumalanga, KwaZulu-Natal and Eastern Cape. It has not been recorded outside of South Africa.

Habitat :

Giant pill-millipedes occur mostly in forest and in densely wooded savanna. They prefer habitats with high moisture and dense leaf litter such as forest floors. They also prefer loam soil because the texture allows infiltration of water and air and is easy for the millipedes to move around in.

Food:

Millipedes are herbivores and they are classified as deposit-feeders, which feed on dead organic matter such as decaying leaves, wood and fruits in the soil and leaf litter.

Life cycle :

Not much is known about the life cycle of pill-millipedes. Some records exist of pill-millipedes living as long as eleven years with females estimated to have produced dozens of batches of eggs during their life span. There are, however, no published studies on the life cycle of the giant pill-millipede and we do not know how long the juveniles take to reach maturity, how long they live for or how long they take to reach their maximum size.

Sex :

Sex organs are situated on the legs closer to the head. The mating behaviour of pill-millipedes can be grouped into four phases. The first phase is when a male detects a female, and orientates itself by positioning its anal shield towards the potential partner. Second, once the male is in contact with the female it starts to make stridulation sounds and vibrations. If the female recognises the male and is receptive, she will open up from her rolled-up position, or not roll up, and the male will then move below the female and grab her front legs with his telopods. The male then ejects sperm from his penises (he has one small penis at the base of each of the first pair of legs on the second segment), and transfers the sperm backwards along his legs and into the female opening which is on her second pair of legs. The two millipedes will then separate. After that, the female lays her eggs in the soil and covers them with a mud layer for protection. The eggs hatch into very small, pale pill-millipedes.

Family life :

Pill-millipedes do not appear to have any form of family life and they live independently. The females do not protect their eggs or young.

THE BIG PICTURE :

Friends and Foes

Giant pill-millipedes are mostly eaten by birds and small mammals such as mongooses and tortoises. They have also been reported to be preyed upon by the carnivorous slug, Chlamydophorus, in the coastal forests of KwaZulu-Natal.

Smart Strategies :

Giant pill-millipedes are terrestrial and mostly active at night. They spend most of the time underground. The burrowing strategy is thought to have evolved as a defence mechanism against predators. They are also capable of rolling into a tight ball, which protects them from predators because it makes them difficult to grasp. Rolling into a ball also enables them to survive in dry conditions.

Poorer world without me :

Pill-millipedes play a major role in the ecosystem by contributing to the transformation of detritus into humus. This process contributes to soil fertility by increasing the availability of nutrients such as nitrogen and phosphates for plant growth.

People & I

Millipedes are harmless to humans. Some of the pill-millipede species have been recorded from traditional medicine markets where they are traded.

Conservation status and what the future holds

Few studies have been conducted on the population status of southern African millipedes. Since the giant pill-millipede is quite habitat specific, the main threats would be loss of habitat through development, especially along the coast. Overharvesting for the traditional medicine trade may be an additional threat. The pill-millipedes have not been assessed according to the IUCN Red List criteria.

Relatives :

Pill-millipedes (Order: Sphaerotheriida) are widely distributed in tropical and sub-tropical environments where more than 200 species are known to occur in Madagascar, Australasia, Southeast Asia and southern Africa. The genusSphaerotherium is endemic to Africa and 54 species have been described from southern Africa. Most of these species are found in coastal areas with the highest number recorded in KwaZulu-Natal. Glomeris is another genus of pill-millipede that occurs in the northern hemisphere, but it is not related toSphaerotherium and it belongs to a different order, Glomerida. The Glomerisspecies are distinct from sphaerotheriids in that they have poison glands that secrete chemicals to repel predators, while the sphaerotheriids do not have such glands.

References and further reading:

Herbert, D.G., Hamer, M.L., Mander, M., Mkhize, N. & Prins, F. 2003. Invertebrate animals as a component of the traditional medicine trade in KwaZulu-Natal, South Africa. African Invertebrates 44,2: 327–344.
Lawrence, R.F. 1987. The biology of the cryptic fauna of forests with special reference to the indigenous forests of South Africa. A.A. Balkema, Cape Town.
Lawrence, R.F. 1953. The centipedes & millipedes of southern Africa. A Guide. A.A. Balkema, Cape Town.
Van Den Spiegel, D., Golovatch, S.I. & Hamer, M. L. 2002. Revision of some of the oldest species in the millipede genusSphaerotherium Brant, 1833 (Diplopoda, Sphaerotheriida, Sphaerotheriidae), with new synonymies. African Invertebrates 43: 143–181.
Wesener, T.,Köhler, J., Fuchs, S. & Spiegel, D. 2011. How to uncoil your partner—‘mating songs’ in giant pill-millipedes (Diplopoda: Sphaerotheriida). Naturwissenschaften, 98,11: 967–975 DOI:10.1007/s00114-011-0850-8.

Some millipedes need “good vibrations” to mate!

courtesy to :www.africamuseum.be/museum/research/general/research-picture/millepedes-good-vibrations

Didier Van den Spiegel (Invertebrates non-insects section) and three German

researchers, Thomas Wesener (Research Museum Alexander Koenig), Jörn Köhler (Hessisches Landesmuseum) and Stefan Fuchs (Goethe Universität), have studied the sounds emitted by certain African millipedes. They have discovered that the female millipedes only open up to the males when they produce sounds, or rather a particular range of vibrations.

Volvation:

The Sphaerotheriida, a group of millipedes, can roll up completely in a ball, just like hedgehogs and pangolins do, in order to protect themselves from predators. However, this phenomenon known as volvation is a major problem at the time of mating. The female rolls up as soon as she is afraid, even if it is a male of her own species that is approaching her.

Vibrations:

In order to mate with her, the male has to “uncoil” the female. To do this he lets out a specific signal by rubbing special ribs on the last pair of legs across nubs on the body shield. The male thus produces sounds similar to those made by cicadas. But millipedes don’t have any acoustic organs. In other words, they’re deaf! So you may well wonder why the males produce sounds at all. In fact the females detect thevibrations emitted by stridulations. Moreover, by studying these complex mating mechanisms, researchers have also discovered that each species produces a different song. So a female only opens up to a male if he is giving out the right vibrations!

History of the research :

This research was begun in 1972 by the Dr. Haacker and Stefan Fuchs, his student at the time. Unfortunately, a few weeks after their return from South Africa, Haacker died of leukaemia. Several decades later, researchers displayed a renewed interest in this study. But the specimens gathered by Haacker could not be identified until 2002, when Didier VandenSpiegel and other colleagues published a taxonomic revision of the giant pill-millipedes from South Africa.

More than 30 years later, the study led to a publication. The article “How to uncoil your partner - "mating songs" in giant pill-millipedes (Diplopoda: Sphaerotheriida)” is published in the Naturwissenschaften review (number 98 (11), pages 967-975) in November 2011:

http://www.springerlink.com/content/u3h307757876t723/

Zoosphaerium solitarium Wesener, 2009, a giant pill-millipede species discovered in 2009 on Madagascar. This species is only known from a single hill in northern Madagascar that is endangered by ongoing forest destruction. Photo: Jörn Köhler.

Unlike other millipedes, giant pill-millipedes cannot excrete poisonous defence fluids. The rolling-in behaviour is a good defence against predators, but leads to communication break-down with a potential mate. Photo: Jörn Köhler.

Sphaerotherium giganteum is the largest species of its genus. Its size can easily reach 6 cm.

Photos: P. Koomen

MEET THE GIANT PILL MILLIPEDE, DOUG.

Figure 1. Organisms like H. haydeniana are found in the Pacific Northwest region of North America.

Giant Pill Millipedes

The Night Train Millipede
The Clown Millipede
The Yellow-Spotted Millipede
The Almond-Scented Millipede
The Cyanide Millipede

courtesy to :bioweb.uwlax.edu/bio203/f2013/crain_alex/

Classification by S u n i t a N a n d i h a l l i:

Domain: Eukaryota
This organism’s cells have membrane-bound nuclei and organelles. (Campbell et al. 2008).
Kingdom: Animalia
This organism is eukaryotic, multicellular, heterotrophic, is motile at some stage in its life cycle, and lacks a structural cell wall. (Campbell et al. 2008).
Phylum: Arthropoda
This organism is an invertebrate that has an exoskeleton and a segmented body with jointed limbs.(Campbell et al. 2008).
Superclass: Myriapoda
This organism is an arthropod whose body is composed of numerous segments each adorned usually with two pairs of legs. (Ruppert & Barnes 1994)
Class: Diplopoda
This organism is a millipede. (Blower 1985)
Order: Polydesmida
This organism is a true, flat-backed millipede. (Hopkin & Reed, 1992).

Family: Xystoesmida
This organism is a millipede that lacks eyes, has stink glands, and has gonopods on the first pair of legs on the seventh segment. (Schimming & Shelley 2008).
Genus: Harpaphe
This organism is a millipede that lives on the Pacific coast from southern Alaska to central California. (Chamberlin & Hoffman, 1958).
Species: Haydeniana
This organism is a millipede with a shiny black body and bright yellow patches on the outside portion of most of its segments. (Schimming & Shelley 2008).

This article dedicated to furthering your knowledge about a millipede called Harpaphe haydeniana.
H. haydeniana is the scientific name, but if you swear you've seen this millipede before, here are some of the many common names that it goes by...

Figure 1. Millipedes including H. haydeniana belong to the phylum Arthropoda

Figure 2. A breakdown of the class Diplopoda shows the location of the order Polydesmida on the phylogenetic tree.

References in Order of Appearance:

Campbell, N.A., Reece, J.B., Urry, L.A., Cain, M.L., Wasserman, S.A., Minorsky, P.V., Jackson, R.B. 2008. Biology. Pearson Benjamin Cummings, San Francisco, California, U.S.A.

Ruppert, E.E. and Barnes, R.D. 1994. Invertebrate Zoology. Saunders College Publishing, Fort Worth, U.S.A.

Blower, J.G. 1985. Millipedes keys and notes for the identification of the species. The Linnean Society of London, London, England.

Hopkin, S.P. and Read, H.J. 1992. The biology of millipedes. Oxford University Press, New York, U.S.A.

Schimming, L. and Shelley, R. 2008. Species harpaphe haydeniana. Iowa State University Entomology. <URL: http://bugguide.net/node/view/15008> Accessed 9 November 2013.

Chamberlin, R.V. and Hoffman, R.L. 1958. Checklist of the millipedes of North America. United States Government Printing Office, Washington, D.C., U.S.A.

Photo References:

Wikimedia Commons 2011. <http://commons.wikimedia.org/wiki/File:Harpaphe_haydeniana_0446.JPG> Accessed 6 December 2013.

Original drawings by Sunita Nandihalli 2013

Habitat and Geography by S u n i t a N a n d i h a l l i

H. haydeniana is found to live in the Pacific Northwest region of the United States. This includes Oregon, Washington, northern California, and Vancouver Island. The type locality (or the area where this species was first identified) of H. haydeniana was Oregon, U.S.A. (Chamberlin & Hoffman 1958).

Since millipedes feed on dead plant material and fragments of organic matter (Hopkin & Reed 1992) litter layer in mature conifer forests with Douglas-fir is H. haydeniana’s habitat of choice. (Acorn & Sheldon 2001). Adults are found in the upper layer, immature stages deeper in the litter. However, the leaf litter in the Pacific Northwest consists of a lot more than just pine trees. Ash, Sycamore, Aspen, Birch, Cedar, Cherry, Chestnut, Cypress, Dogwood, Fir, Elm, Oak, Maple, Spruce, and Redwood are just a few more common trees that grow in this area and that H. haydeniana has fed on as well. (Jensen et al. 2011).

It has been shown in numerous cases that millipedes are better for plant growth and, in turn, ecosystems altogether, than even worms. This is because H. haydeniana excretes decomposed organic matter in the form of fecal pellets. (Thakur et al. 2011). It was found that higher values of Nitrogen, Phosphorous, Potassium, Magnesium, and Calcium (necessary nutrients in order for plants to produce enzymes and nucleic acids) were observed in millipede compost as opposed to worm compost or ordinary soil. Since millipede feces are so high in nutrients, it is not uncommon for them to partake in coprophagy or the eating of one’s own feces in order to gain extra nutrients that you previously didn’t during the first full cycle of digestion.

Although the chemical defenses of H. haydeniana are able to keep most predators at bay, one specific beetle is specialized in consuming H. haydeniana. P. laevissimus also lives in California, Oregon, and Washington and feeds on millipedes like H. haydeniana. (Denton 1997).

Some other organisms that live in the Pacific Northwest Region of the United States include T. longiglossus, D. maculata, H. thysbe, C. felis, and V. vinifera.

Figure 2. The Bald-faced hornet, D. maculata, is another arthropod that lives in the Pacific Northwest

References in Order of Appearance:

Chamberlin, R.V. and Hoffman, R.L. 1958. Checklist of the millipedes of North America. United States Government Printing Office, Washington, D.C., U.S.A.

Hopkin, S.P. and Read, H.J. 1992. The biology of millipedes. Oxford University Press, New York, U.S.A.

Acorn, J. and Sheldon, I. 2001. Bugs of Washington and Oregon. Lone Pine Publishing, Edmonton, Canada.

Jensen, E., Zahler, D., Patterson, B., & Littlefield, B. 2011. Common trees of the pacific northwest. Oregon State University. <URL: http://oregonstate.edu/trees/name_common.html> Accessed 9 November 2013.

Thakur, P.C., Apurva, P., and Sinha, S.K. 2011. Comparative study of characteristics of biocompost produced by millipedes and earthworms. Advances in Applied Science Research 2: 94-98.

Denton, M. 1997. Promecognathus laevissimus dejean 1829. The Evergreen State College. <URL: http://academic.evergreen.edu/projects/evergreenBiota/kingdom/animalia/phylum/arthropoda/class/insecta/order/coleoptera/family/carabidae/
GENERA/PROMECOGNATHUS/P_laevissimus.HTM > Accessed 9 November 2013.

Photo References in Order of Appearance:

Wikimedia Commons 2012. <http://commons.wikimedia.org/wiki/File:Pacific_northwest-relief.png> Accessed 19 November 2013.

Wikimedia Commons 2012. <http://commons.wikimedia.org/wiki/File:July_31,_2012_-_Bald-faced_Hornet_on_Red_Flower.jpg> Accessed 5 December 2013.

Morphology by S u n i t a N a n d i h a l l i

External Morphology:

The morphology of Harpaphe haydeniana and millipedes in general consist of the head, body, and telson. (Hopkin & Reed 1992). The head itself is calcified heavily in order to facilitate burrowing in leaf litter where these millipedes live. The head consists of the mouth, and sensory organs. The mouth has two parts, the mandibles and the first maxillae. The mandibles are for biting and crushing and the maxillae are used for tasting and also chewing. The head also bears sensory organs such as the antennae and Tӧmӧsváry organs. Since H. haydeniana possesses no eyes and is therefore blind, it used its antennae to feel and sense the surrounding environment. The exact function of the Tӧmӧsváry organ is unknown, but it is strongly suggested due to numerous studies that have been done that these organs are receptive to sights, smells, and sounds. (Hopkin & Reed 1992).

Figure 1. H. haydeniana mingling about the leaf litter

H. haydeniana’s body is about 1.8 inches in length (Acorn & Sheldon, 2001) and has 20 twenty segments (Hopkin & Reed 1992). The surface of the body of H. haydeniana is smooth, black, and has bright yellow spots that run along the millipede’s side. This bright coloring is a warning to any predator that it is poisonous, as it secretes cyanide which is a powerful poison. (Acorn & Sheldon 2001). Although H. haydeniana is known for its chemical defenses and bright coloring, millipedes in general are mostly known for its legs. The word millipede comes from name meaning “thousand foot”. (Acorn and Sheldon 2001). The legs of millipedes grow ventrally which requires the S-shaped structure that they have, giving them the appearance of “hanging down” from their legs rather than standing on them. (Hopkin & Reed 1992). Since H. haydeniana has so many legs (two pairs per segment), they run the risk of running into each other. To combat this, H. haydeniana moves its legs in slow, coordinated, metachronal waves that start at the back of the body and move towards the head. (Acorn & Sheldon 2001). Males tend to have longer legs than females in order to more strongly grasp the females during copulation. Besides this difference between sexes, each leg is the same length throughout the body. The cuticle of each segment of millipedes consists of a dorsal tergite, ventral sternite, and lateral pleurites. In polydesmid millipedes like H. haydeniana, each segment is strengthened by the fusion of all three of these structures. (Hopkin & Reed 1992).

Figure 2. The head consists of organs such as the antenna and mandibles, the body consists of tergites, sternites, and legs. The body of H. haydeniana ends with the telson.

The cuticle itself consists of the three layers, the very thin epicuticle, exocuticle, and endocuticle. H. haydeniana, as well as most other millipede species, have a calcified cuticle that they accumulate when they eat decaying organic material. It is also permeable to water, restricting this millipede’s habitat to humid areas. Since H. haydeniana is an arthropod, it has an exoskeleton and must do ecdysis. The timing of this molting process is under hormonal control. (Hopkin & Reed 1992). On the ventral side of body are the reproductive structures. The gonopods of adult male millipedes are located on the seventh segment and replace one or both legs. Female gonopods are internal but may be extruded during copulation. When this happens, they can be seen behind the second pair of legs. Each oviduct of female millipedes opens separately into organs called vulvae which are in separate sacs within the lumen and these are the structures that are everted during copulation. (Hopkin & Reed 1992).

The telson which is the last division of the body consist of a pre-anal segment, a pair of anal plates, and a sub-anal scale. The anal plates form a valve that opens during defecation. (Blower 1985). 80 to 90 percent of dry food ingested by H. haydeniana is excreted as feces. The two forms of predominating nitrogenous waste in millipedes are ammonia and uric acid. Ammonia must be excreted quickly in order to avoid self-poisoning but uric acid can be stored temporarily in the midgut epithelium. (Hopkin & Reed 1992).

Internal Morphology :

The digestive tract of a millipede is basically a straight tube from mouth to anus. Small pieces of dead plant material are passed into the lumen of the foregut where it receives secretions from salivary glands to moisten the food. Actual digestion takes place in the midgut, where enzymes secreted by epithelial cells break down the plant material into its simple chemical compounds. The most important site in any millipede for assimilation of nutrients is the midgut epithelium. Products of digestion are absorbed by the microvilli that border the cells in the midgut and are then intracellularly digested and passed to the liver which is a structure that is a layer of cells that surround the midgut. (Hopkin & Reed 1992). Adipocytes or lipocytes, fat cells that primarily compose of adipose tissue (specialized in storing energy as fat), serve as part of the digestive system. They function with the excretory system. (Camatini 1979). The organs involved in the excretory process include the midgut epithelium, liver, integument, exocrine glands, haemocytes, nephridial organs, nephrocytes, ecdysial glands, malpighian tubules, and the fat body. Millipedes have one pair of true excretory organs called nephridial organs and regulate excretion in these organisms. Nephrocytes are cells that take up substances in the haemolymph and which are then metabolized. Some of these products are then stored or returned to the circulation.

Millipedes, being arthropods, have an open circulatory system. This means that the blood of the animal, composed of liquid and cellular components are circulated throughout the body via the pumping action of the heart or dorsal vessel. The liquid bathes the organs in oxygen and nutrients and transports the products of metabolism to and from organs of digestion, storage, and excretion. This fluid in the body cavity is called the hemocoel. There is no distinction between blood and interstitial fluid; this combined fluid is called hemolymph. (Campbell et al. 2008). The principle sugar is trechalose and the main lipids are phospholipids in the hemolymph. The blood of millipedes also transports nitrogeneous wastes in a form that can be tolerated and can be excreted. (Camatini 1979).

References in Order of Appearance:

Hopkin, S.P. and Read, H.J. 1992. The biology of millipedes. Oxford University Press, New York, U.S.A.

Acorn, J. and Sheldon, I. 2001. Bugs of Washington and Oregon. Lone Pine Publishing, Edmonton, Canada.

Blower, J.G. 1985. Millipedes Keys and Notes for the Identification of the Species. The Linnean Society of London, London, England.

Camatini, M. 1979. Myriapod biology. Academic Press Inc., London, Great Britain.

Campbell, N.A., Reece, J.B., Urry, L.A., Cain, M.L., Wasserman, S.A., Minorsky, P.V., Jackson, R.B. 2008. Biology. Pearson Benjamin Cummings, San Francisco, California, U.S.A.

Photo Citations:

Original drawing by Sunita Nandihalli 2013.

Wikimedia Commons 2012. <http://commons.wikimedia.org/wiki/File:Harpaphe_haydeniana_0447.JPG> Accessed 20 November 2013.

Figure 1. Maple leaf litter would be an ideal meal for a millipede

Nutrition by Alex Crain

The H. haydeniana prefers dead organic material, such as leaf litter; however they will eat dead arthropods as well as roots and the shoots of young seedlings. Millipedes have also been known to consume their own feces. (Hopkin & Read 1992) The millipede’s mouth consists of a pair of maxillae, a pair of mandibles and a plate known as the gnathocilarium. The maxillae acts much like our upper jaw and the mandibles like our lower jaw; the gnathocilarium is literally used as a plate which the mandibles works against when shredding food like leaf litter. (Branson 1997)

However H. haydeniana is not the organism that is considered a decomposer, some other examples are L. edodes, A. alternata, G. tappaniana, A. muscaria and G. sterkii

In the rainforests on the coasts of British Columbia millipedes are the main macrofauna and consume a large proportion of the annual litter, H. haydeniana alone consumes close to 36% of the litter. (Cárcamo et al. 2000) Although in some of the millipede’s food sources, especially leaf litter there are other chemicals that are not necessarily nutrients that have dietary effects on the millipede, mostly negative. Phenol and tannic acid are some of these detrimental chemicals, which are naturally occurring within plants in small quantities. In an experiment where the millipedes are exposed to less than 10% of the acid for 7 days showed a statistically significant decrease in rate of assimilation, as well as growth rate in the younger subjects. There was also evidence of lower level of tissue glycogen, carbohydrates and lipids. (Roy and Joy, 2009) However it was shown that the millipedes are very sensitive to these chemical amounts and will feed on certain leaf litter types if given a choice. (Joy et al. 1998)

Figure 2. H. haydeniana crawling though its favorite food, leaf liter

The digestion of the litter takes place primarily in the foregut of the millipede where the salivary glands secrete enzymes, including catalase, cellulose and peroxidase. (Hartenstein 1982) After completely passing through the gut, undigested materials are released in the form of fecal pellets. About 90% of leaf litter, by mass, consumed by the millipede is egested in this form. (David & Gillon 2002) A comparison of the chemical composition of leaf litter versus the millipede’s feces indicates that a millipede utilizes only water-soluble and/or compounds such as carbohydrates, short-chained amino acids and lipids, compounds that are easily degradable. (Rawlins et al. 2006)

All millipedes are oviparous, which means that they lay eggs. The most common place for a H. haydeniana millipede to lay her eggs is under a rotten log. There have been some cases among millipedes where the male millipede participates in the act of egg-guarding, standing guard while the eggs are developing. (Requena et al. 2009). Millipedes lay somewhere between 30 and 300 eggs each breeding season. The young hatch after three or more weeks of incubation, sometime during the late spring. (Branson 1997) Reproducing can be quite taxing on the female, according to a study done at the Kansas Academy of Science. It was shown that in some species the more often a female reproduces the shorter her life span. Individual females collected in April mated about 5 times each compared to individuals who were collected in March, who only mated an average of 1.75 each. April individuals lived about a month, whereas March individuals lived over 4. ( Youngsteadt 2009)

Reproduction by Alex Crain

The mating season for the millipede H. haydeniana occurs in the spring. During the mating season the male and female millipede come towards each other head on, where they meet and then raise their anterior ends. Next they climb up each other’s ventral side and a female millipede receives sperm from the male, which is stored in the seminal receptacles located in the vulvae. (Blower 1985) Both the female and the male’s reproductive openings are located on the third body segment, just behind the second pair of legs. The female’s is surrounded by two different plates, one of which

covers the opening. (Branson 1997) The H. haydeniana male does what is called direct sperm transfer, where the sperm is placed directly into the female opening. This transfer is made possible through the use of special intromittent organs known as gonopods. (Blower 1985)

Figure 1. two H. haydeniana during mating season

Figure 2. A fallen log as an ideal spot for a female to lay her eggs

The eggs then hatch into an immobile pupoid, which then molts into an instar. At the instar stage, the millipede has four developed rings (segments) and three pairs of legs between the rings. Behind the mobile rings are two to three immobile rings which are then followed by the telson or posterior-most segment of the body, like a tail. By the next molt the immobile rings will all have pairs of legs. (Blower 1985)

Figure 3. The general anterior anatomy of a millipede

Within millipedes there are three types of anamorphosis or a change to a higher form. The first type being euanamorphosis, where each molt is accompanied by the addition of a new ring, even after the millipede has reached sexual maturity. The second type is called teloanamorphosis, where the addition of rings stops at a certain stage and the molting stops. Finally the third type is known ashemianamorphosis where the addition of new rings continues until a certain stage, however molting continues. (Blower et al. 1993) H. haydeniana exhibits the second type of anamorphosis orteloanamorphosis, their molting stops after they have reached sexual maturity. (Blower 1985)

References in Order of Appearance:

Blower, J.G. 1985. Millipedes: Keys and Notes for the Identification of the Especies. The Bath Press, Avon, UK.

Branson, B. A. 1997. Legions of legs. World & I 12: 170.

Requena, S.R. et al. 2009. Efficiency of uniparental male and female care against egg predators in two closely related syntopic harvestmen. Animal Behavior 78:1169-1176

Youngsteadt, N.W. 2009. Laboratory Observations on the Natural History of Pseudopolydesmus pinetorum (Diplopoda, Poydesmida, Polydesmidae) with Emphasis on Reproduction and Growth. Transactions of the Kansas Academy of Science 112: 67-76.

Blower, J.G., Dohle W. and Enghoff H. 1993. Anamorphosis in millipedes (Diplopoda)- the present state of knowledge with some developmental and phylogenetic considerations. Zoological Journal of the Linnean Society 109: 103-234

Photos References in Order of Appearance:

Wikimedia Commons 2011.<http://commons.wikimedia.org/wiki/File:Harpaphe_haydeniana_043.jpg> Accessed 6 December 2013.

Wikimedia Commons 2009.<http://commons.wikimedia.org/wiki/File:Fallen_Log_(3728017912).jpg> Accessed 6 December 2013.

Wikimedia Commons 2013.<http://commons.wikimedia.org/wiki/File:Millipede_anterior_anatomy.png> Accessed 6 December 2013.

Adaptations & Interactions

Adaptations by Alex Crain

The H. haydeniana has a unique defense mechanism when it comes to deterring predators from eating them, they have the ability to produce the toxin, hydrogen cyanide (HCN).
When the yellow spotted millipede discharge hydrogen cyanide, the secretion oozes from the glands onto the millipede’s surface and coats the exoskeleton of the animal. On the sides of the millipede’s body are two-part glands that consist of a reservoir, which stores cyanohydrin and a reaction chamber where the cyanohydrin is broken down by enzymes to create hydrogen cyanide; as well as an aromatic molecule, benzaldehyde. The benzaldehyde is what gives the millipede’s their distinctive “bitter almond” smell. (Eisner et al. 1963) The millipede also metabolizes small quantities of the cyanide into β-cyanoalanine as well as asparagine; these two compounds may be used to detoxify small amounts of HCN that may leak into the milliped's body from the storage chamber housing the toxic chemical. (Rockstein 1978)..

The millipede's secretions act as a topical irritant. As a general rule millipedes eject their toxins during the beginning stages of an attack, deterring their adversary before receiving major injuries. (Eisner and Meinwald 1966) Millipedes in contrast to centipedes do not have fangs, however the toxins that they secrete can cause erythema and brown/black pigmentation of the affected area.

Figure 1. This is what happens when toes are exposed to H. haydeniana's toxin

The pigmented lesions may last as long as several months. Though the effect of millipede toxins may look serious (on the skin) there are no major repercussions in humans. It is advised to immediately wash the effected site with alcohol or ether in order to dissolve the toxins. However if one were to get the toxin into their eye, seek immediate help from an ophthalmologist; in order to prevent blindness. (Hudson and Parsons 1997)

Interactions by Sunita Nandihalli

H. haydeniana has both positive and negative interactions with other organisms. A negative interaction would be any interaction that either has a negative effect on the millipede or other organisms. A positive interaction would be a relationship (direct or indirect) that would have a positive effect on the millipede or the millipede and other organisms.

Let’s start out with negative organismal interactions: parasites and predators of H. haydeniana. Nematodes are one of the most common types of endoparasite among millipedes along with Dipterans (flies) in the family Phaeomyidae. Beetles, reptiles, birds, shrews, raccoons and other various mammals, spiders, amphbians, and even snails have been documented to prey on millipedes. (Sierwald & Bond 2007). However, when these animals get a taste for hydrogen cyanide, they’re most likely not going to want to get anywhere near H. haydeniana again. Although these chemical defenses of H. haydeniana are able to keep most predators at bay, one specific beetle is specialized in consuming H. haydeniana. P. laevissimus also lives in California, Oregon, and Washington and feeds on millipedes like H. haydeniana. (Denton 1997).

Figure 1. Raccoons are one of the many diverse organisms that may feed on millipedes.

Not all H. haydeniana interactions are negative, however, some inter-organismal relationships are quite beneficial. For example, Trichomycetes are obligatory symbiotrophic fungi that live in the intestines of millipedes. The fungi’s mycelia help break down the dead plant material while the millipede’s hindgut provides a habitat for the fungi. (Sierwald & Bond 2007).

Interactions References in Order of Appearance:

Sierwald, P. and Bond, J.E. 2007. Current status of the myriapod class diplopoda (millipedes): taxonomic diversity and phylogeny. The Annual Review of Entomology. <URL: http://www.annualreviews.org/doi/pdf/10.1146/annurev.ento.52.111805.090210> Accessed 19 November 2013.

Denton, M. 1997. Promecognathus laevissimus dejean 1829. The Evergreen State College. <URL: http://academic.evergreen.edu/projects/evergreenBiota/kingdom/animalia/phylum/arthropoda/class/insecta/order/coleoptera/family/
carabidae/GENERA/PROMECOGNATHUS/P_laevissimus.HTM > Accessed 9 November 2013.

Interactions Photo Reference:

Wikimedia Commons 2009. < http://commons.wikimedia.org/wiki/File:Raccoon_(4152678243).jpg> Accessed 5 December 2013.

Adaptations References in Order of Appearance:

Eisner, T., H.E. Eisner, J.J. Hurst, F.C. Kafatos, and J. Meinwald. 1963. Cyanogenic glandular apparatus of a millipede.Science 139: 1218-1220

Rockstein, M. 1978. Biochemistry of Insects. Academic Press, New York, New York, USA.

Eisner, T. and J. Meinwald. 1966. Defensive Secretions of Arthropods. Science 153

Hudson, B.J. and G.A. Parsons. 1997. Giant millipede “burns” and the eye. Trans R Soc Trop Med Hyg 91: 183-185

Adaptations Photo References in Order of Appearance:

Wikimedia Commons 2006.<http://commons.wikimedia.org/wiki/File:Millipede_toes_toxin_diplopoda.png> Accessed 6 December 2013.

Refrences :

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Two other species of Harpaphe (H. pottera and H. telodonta) occur within the range of H. haydeniana, both with yellow-tipped paranota. H. telodonta(Humboldt and Del Norte counties, California) is slightly more brown in colour and has more strongly pointed keels while H. pottera (Mendocino and Humboldt counties) can only be distinguished by close examination of the male reproductive organs (gonopods).

The genus Harpaphe is in the family Xystodesmidae, which contains several other species with similar markings, including North American species ofBoraria, Chonaphe, Paimokia, Hybaphe and Montaphe.[4] Exact species determination requires examination of the male gonopods, but the sharply pointed posterior corners of the paranota can help distinguish Harpaphe fromHybaphe and Chonaphe.

Outside of North America, superficially similar species include Anoplodesmus saussurii which has been mistakenly called H. haydeniana,[6] and Asiomorpha coarctata (Paradoxosomatidae), the latter species being native to Southeast Asiabut widely introduced around the world, including the American Gulf Coastregion.[7] The ability to secrete hydrogen cyanide is shared by other members of the Polydesmida, the largest order of millipedes.

Asiomorpha coarctata (Paradoxosomatidae)

BandfüßerTHAIORANGE

Armored millipede (Polydesmidae)

Brachydesmus

Polydesmid millipede

Polydesmus collaris C.L. Koch, 1847