Motyxia

Adult Motyxia reach 3 to 4 cm in length, 4.5 to 8 mm wide, with 20 body segments, excluding the head. Females are slightly larger than males. Like other polydesmidans ("flat-backed" millipedes) they lack eyes and have prominent paranota (lateral keels). They are typically tan to orange-pink in color (except M. pior), with a dark mid-dorsal line. M. pior is the most variable in color, and ranges from dark gray to greenish-yellow to bright orange. They lack bumps on the metatergites (the dorsal plates possessing paranota), giving a somewhat smooth appearance.[2] The anterior 2–3 diplosegments are oriented cephalically (towards the head), a trait most distinct in M. sequoiae, nearly indistinct in Motyxia porrecta. They are fluorescent under black light (millipedes in the tribe Xystocheirini display some of the brightest fluorescence of the U.S. Xystodesmidae species). Most uniquely they are bioluminescent: emitting light of their own production.[1]

An adult M. sequoiae photographed in normal light (A) and its own light (B)

There are 11 species of Motyxia.[3] They some of the few known bioluminescent species of millipedes, a class of about 12,000 known species.[4] Motyxia sequoiae glows the brightest and Motyxia pior the dimmest.[5] Light is emitted from the exoskeleton of the millipede continuously, with peak wavelength of 495 nm (the light intensifies when the millipede is handled).[6] Emission of light is uniform across the exoskeleton, and all the appendages (legs, antennae) and body rings emit light. The internal organs and viscera do not emit light. Luminescence is generated by a biochemical process in the millipede's exoskeleton.[6] The light originates by way of a photoprotein, which differs from the photogenic molecule luciferase in firefly beetles.[7]

Motyxia's photoprotein contains a porphyrin and is about 104 kDa in size.[7][8] However, the structure of the luminescent photoprotein remains uncertain, and its homology to molecules of closely related arthropods is unknown.

Besides Motyxia, the only known bioluminecscent millipedes are Paraspirobolus lucifugus (Spirobolellidae), from Japan and Taiwan, and Salpidobolus (Rhinocricidae), both in the order Spirobolida.[9] Another luminescent species of dubious existence from North America has been claimed, but it has not been found since the 1890s and the luminescence attributed to it is believed to have been larval phengodid beetles, also known as glowworms.[1]

The discovery of the Motyxia bistipita after nearly 50 years revealed new characteristics about the species and genus. In 1967, two small millipedes were discovered in San Luis Obispo, and were coined as a new species, Xystocheir bistipita. For almost 50 years, these two millipedes were the only sightings of the species, until they were rediscovered in 2013 in San Luis Obispo once again. The rediscovered specimens were brought to a lab and examined, where it was discovered that the species was bioluminescent. With this new piece of information, the Xystocheir bistipita was found to have a closer phylogenetic relationship with the Motyxia species and was renamed Motyxia bistipita.[12]

M. bistipita lives in low elevations of the Sierra Nevada mountains in California, in a hotter, drier climate than other Motyxia. The glow is believed to be a response to heat; the bioluminescent proteins help neutralize the body's byproducts caused by heat. The bioluminescence later evolved as a warning signal to predators that the body contained cyanide.[13][12]

An open meadow surrounded by conifers in Sequoia National Park

Motyxias occur in live oak and giant sequoia forests, and notably also in meadows. The presence of xystodesmid millipedes in meadows is atypical for the family. Most species are observed under canopies of broad-leaf deciduous forests. All Motyxia species are exclusively nocturnal. During the day, individuals are burrowed beneath the soil. At night, they emerge (by an unknown mechanism not related to light since they're blind) and feed on decaying vegetation. Individuals of the species M. sequoiae have been observed climbing on tree trunks, possibly consuming algae and lichens adhering to the bark.

Natural predators of motyxias include rodents,[5] geophilid centipedes, and larval phengodid beetles.[1]

The life cycle of M. sequoiae was studied in detail in the early 1950s.[2] Eggs are round, about 0.7mm in diameter, and are laid in masses of 70–160 eggs. After about two weeks the larvae hatch with seven body segments and three pairs of legs, and measure about 1.7mm long. Juveniles exhibit bioluminescence as early as hatching. Additional legs and body segments develop with each molt, during which the animals construct a protective spherical cocoon or molt chamber out of soil. Larvae go through seven developmental stages (instars) before reaching adulthood. In males, the single pair of reproductive structures (gonopods) begin to develop in the 4th instar, before which male and females have equal numbers of walking legs, and after which males have one fewer pair.[2]

Distribution of Motyxia species, after Shelley, 1997[14]

Motyxia species occur in an approximate 280 km (175 mi) vertical range across three counties in California: Los Angeles, Kern and Tulare counties.[1] Species predominately occur in the Santa Monica, Tehachapi, and southern Sierra Nevada Mountains. The northernmost species is M. pior, which occurs as far north as Crystal Cave in Sequoia National Park. The southernmost species is M. monica, which has a disjunct distribution: a population in southern Kern County, and an isolated one in the Santa Monica Mountains near the city of Los Angeles. The range of the eight species largely do not overlap, although M. tularea overlaps with M. kerna and M. sequoiae.[14]

List of Species [15]

The speciation of Motyxia is believed to have been driven by geological events and a drying climate following the most recent Pleistocene glaciation, while reproductive isolating mechanisms include rivers which are at their fullest levels in times when adults are most active, and marked differences in rainfall and suitable habitat across mountainous terrain.[1]

Motyxia is a member of the family Xystodesmidae, a group of large and colorful millipedes, and M. monica is the southernmost species of Xystodesmidae in western North America.[1] Within Xystodesmidae, Motyxia is placed in the tribe Xystocheirini along with the genera Anombrocheir, Parcipromus, Wamokia, and Xystocheir, all of which are native to California.[16]

According to Paul E. Marek, one of the most heavily involved researchers of these millipedes, "much about Motyxia remains mysterious." There is still a lot to learn about Motyxia's mating habits, their emergences into the night, and the exact evolutionary reasons for their bioluminescence.[17]

• J. Rosenberg; V.B. Meyer-Rochow (2009). "Luminescent myriapoda: a brief review". In V.B. Meyer-Rochow (ed.). Bioluminescence in Focus: A Collection of Illuminating Essays. pp. 139–147. ISBN 978-81-308-0357-9.

• Glowing Millipedes: National Public Radio, Science Friday
• Strange Glowing Millipedes Ooze Cyanide to Foil Predators: LiveScience
• Evolution of bioluminescence in millipedes: apheloria.org