Cretoperipatus burmiticus is an extinct species of peripatid velvet worm known from multiple specimens in Burmese amber. It lived in Myanmar's Kachin State during the Cenomanian Age of the Late Cretaceous. Some fossils were exceptionally preserved, allowing detailed analysis of its fine anatomy. Cretoperipatus had a pair of antennae, two simple eyes, clawed legs, and a pair of slime papillae, showing it was it was a crown-group onychophoran. Based on details of it's fine anatomy, Cretoperipatus was placed in the family Peripatidae, likely grouping with basal Asian species like Eoperipatus and Typhloperipatus.
| Cretoperipatus Temporal range: Cenomanian-Turonian
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Scientific classification 
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Onychophora |
| Family: | Peripatidae |
| Genus: | †Cretoperipatus Engel & Grimaldi, 2002 |
| Species: | †C. burmiticus
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| Binomial name | |
| †Cretoperipatus burmiticus | |
Discovery and naming
editBoth the holotype (AMNH Bu218) and various topotypes (BU-001467, BU-001468, ZZZ0066) were found in Burmese amber near Tanai in Myanmar's Kachin State.[1][2] Additional unsubscribed specimens were found in the same location, showing this animal was fairly abundant.[3] All fossils date to around 100 million years ago, meaning the animal lived at the beginning of the Late Cretaceous during the Cenomanian Age.[2][3]
The genus name "Cretoperipatus" is derived from "creto-", since it lived in the Cretaceous and "Peripatus", the type genus of the family Peripatidae. The species name "burmiticus" references the fossil being encased in Burmese amber.[1]
Description
editDue to being encased in amber, many Cretoperipatus specimens preserve minute details that would otherwise disappear in the typical fossilization process. For instance, some fossils preserve scale ranks, or rows of tiny protrusions on the animal’s dermal papillae.[2]
Head
editCretoperipatus had a single pair of antennae with simple eyes called ocelli at their base. On these antennae were multiple rings of specialized, spindle-shaped papillae known as sensilla.[2] Cretoperipatus had a ventral (bottom-facing) mouth with two internal jaws. In the original description, the animal’s genital pad was mistaken for the mouth.[1][2] This was due to the holotype's poor preservation and the two structures looking similar.[2] In addition to the features mentioned above, Cretoperipatus also had a pair of slime papillae. These were somewhat hard to distinguish, even when using x-ray microscopy.[2]
Trunk
editCretoperipatus has around twelve annuli or rings of papillae on each segment of its trunk.[1] Most of these were primary papillae which provide the animal a sense of touch. The primary papillae are divisible into three parts: a larger part on the bottom known as a basal piece, a smaller part in the middle known as the apical piece, and a bristle on top used for sensing the environment. In Cretoperipatus, the basal piece is easily distinguishable from the apical piece and has five scale ranks. The apical piece is asymmetrically shaped due to having three scale ranks in the front and two in the back.[2] In addition to the primary papillae, the trunk also possessed crater-shaped accessory papillae with no bristle.[2] At the very end of its trunk, Cretoperipatus had a genital pad between its penultimate (second to last) pair of legs.[2]
Lobopods
editCretoperipatus had a pair of legs known as lobopods on each segment of its trunk. Like in other velvet worms, the lobopods had a papillae-covered base, spinose pads or cushions, and a clawed foot.[1][2] The animal had no coxal organs on any legs and lacked crural papillae on the front-most pairs. However, all legs have a nephridal opening at their base.[1]
Cretoperipatus was originally interpreted as having only three spiny pads per lobopod,[1] yet better specimens show most of them have four.[2] The original describers probably mistook the holotype's back with its front as this specimen was poorly preserved, and a velvet worm’s endmost legs have fewer cushions.[2] On a typical leg of Cretoperipatus, the distal (furthest away from the trunk) and proximal (closest to the trunk) cushions are thinner than the two between. The third cushion (2nd closest to the body) was also split in two by a nephridial tubercle, an organ used to dispel waste.[2]
Cretoperipatus’s foot had five basal papillae, with the bottom-most two being weakly developed.[1] At the end of the foot were two distal papillae. These were located on either side of a pair of claws.[2] Many of these claws were detached and found floating in the amber matrix. This makes sense, as in modern velvet worms, the claws are connected by only a fragile membrane of tissue.[2]
Classification
editCretoperipatus burmiticus is one of the two fossils confidently assigned to Onychophora, the other being the Late Carboniferous Antennipatus.[3][4] Based on its morphology, Cretoperipatus was assigned to the family Peripatidae. It groups with basal, Southeast Asian species like Eoperipatus totoro and Typhloperipatus williamsoni.[2] Cretoperipatus is likely the closest relative of Typhloperipatus, however, this has not been phylogenetically tested.[3]
Cretoperipatus was assigned to Peripatidae based on the following characteristics: the papillae on its back could be divided into two parts (basal and apical), its genital pad was between the penultimate legs, it possessed spindle-shaped sensilla on its antennae, and it had a diastema on its jaw blade.[2]
Cretoperipatus was grouped with various South Asian species due to having two distal papillae (other peripatids have three or more). It lacked ventral fields of modified scales present in Eoperipatus totoro, but these are also lacking in other species of Eoperipatus. Unlike Eoperipatus but similar to Typhloperipatus, the foot's nephridial tubercle was on the third spinous pad rather than higher up. However, unlike Typhloperipatus, Cretoperipatus still had eyes.[2]
Implications
editZoogeography
editEarly on, it was hypothesized that onychophorans arrived in Asia due to northwards drift of India, which, at the time, was an independent continent. This was known as the “Out of India” hypothesis and explained how velvet worms colonized South and Southeast Asia.[2] Cretoperipatus refutes this, as it had clear affinities with Asian velvet worms and was present before the Asia-India collision during the Late Oligocene. Some time after this, peripatids began colonizing Northeast India, probably from Myanmar.[2] Combined with the presence of other Northern Hemisphere onychophorans (Antennipatus, Helenodora, and tentatively Succinipatopsis), velvet worms occurred in Laurasia (the precursor of North America and Eurasia) far longer than originally thought.[2][5]
References
edit- ^ a b c d e f g h Grimaldi, David A.; Engel, Michael S.; Nascimbene, Paul C. (March 2002). "Fossiliferous Cretaceous Amber from Myanmar (Burma): Its Rediscovery, Biotic Diversity, and Paleontological Significance". American Museum Novitates (3361): 1–71. doi:10.1206/0003-0082(2002)361<0001:FCAFMB>2.0.CO;2. hdl:2246/2914. S2CID 53645124.
- ^ a b c d e f g h i j k l m n o p q r s t u v w Oliveira, I. S.; Bai, M; Jahn, H; Gross, V; Martin, C; Hammel, J. U.; Zhang, W; Mayer, G (2016). "Earliest Onychophoran in Amber Reveals Gondwanan Migration Patterns". Current Biology. 26 (19): 2594–2601. Bibcode:2016CBio...26.2594O. doi:10.1016/j.cub.2016.07.023. PMID 27693140.
- ^ a b c d Giribet, Gonzalo; Buckman-Young, Rebecca S.; Costa, Cristiano Sampaio; Baker, Caitlin M.; Benavides, Ligia R.; Branstetter, Michael G.; Daniels, Savel R.; Pinto-da-Rocha, Ricardo (2018). "The 'Peripatos' in Eurogondwana? – Lack of evidence that south-east Asian onychophorans walked through Europe". Invertebrate Systematics. 32 (4): 840–863. doi:10.1071/IS18007.
- ^ Garwood, Russell J.; Edgecombe, Gregory D.; Charbonnier, Sylvain; Chabard, Dominique; Sotty, Daniel; Giribet, Gonzalo (2016). "Carboniferous Onychophora from Montceau-les-Mines, France, and onychophoran terrestrialization". Invertebrate Biology. 135 (3): 179–190. doi:10.1111/ivb.12130. ISSN 1077-8306. PMC 5042098. PMID 27708504.
- ^ Murienne, Jerome; Daniels, Savel R.; Buckley, Thomas R.; Mayer, Georg; Giribet, Gonzalo (2014-01-22). "A living fossil tale of Pangaean biogeography". Proceedings of the Royal Society B: Biological Sciences. 281 (1775): 20132648. doi:10.1098/rspb.2013.2648. PMC 3866409. PMID 24285200.
External links
edit
Data related to Cretoperipatus at Wikispecies
