TL;DR
Fossil evidence shows that early ancestors of millipedes and centipedes already had legs adapted for underwater environments. This challenges previous ideas that such limbs evolved solely for terrestrial life. The findings shed light on the evolution of myriapods and their transition from water to land.
Fossil evidence from 437 million years ago shows that an ancestor of modern millipedes and centipedes already possessed legs adapted for underwater life, challenging previous assumptions about limb evolution in myriapods.
The fossils, found in the Silurian Brandon Bridge Formation in Waukesha, Wisconsin, include 35 well-preserved specimens of a newly identified arthropod named Waukartus muscularis. These fossils reveal that Waukartus had many unbranched, uniramous legs, with at least 11 segments, and features multiple head appendages likely used for sensory or feeding functions.
Phylogenetic analysis places Waukartus just outside the crown group of myriapods, which have at least 17 limb pairs. Its features suggest it was aquatic, with limbs adapted for underwater movement, yet it possessed limb traits—such as uniramous limbs—that are now associated with terrestrial adaptation. This indicates that limb reduction in myriapods occurred before they transitioned onto land, representing an exaptation rather than a direct adaptation for terrestrial life.
Why It Matters
This discovery rewrites parts of the evolutionary history of myriapods, showing that their ancestors developed leg structures suited for aquatic environments well before moving onto land. It suggests that limb reduction and specialization were not solely driven by terrestrial adaptation but may have involved pre-adaptations that facilitated the eventual terrestrial transition. This insight enhances understanding of arthropod evolution and the complex pathways leading to terrestrial colonization.

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Background
Prior to this discovery, the fossil record of early myriapod relatives was sparse, with limited evidence regarding their aquatic or terrestrial origins. The identification of Waukartus muscularis fills a gap by providing direct fossil evidence of an aquatic ancestor with limbs already showing some terrestrial traits. Earlier studies identified euthycarcinoids as stem-group myriapods, but Waukartus offers a closer look at limb evolution and habitat transition in this lineage.
“Waukartus muscularis provides compelling evidence that the origins of limb reduction in myriapods predate their move onto land, indicating a more complex evolutionary pathway than previously thought.”
— Dr. Jane Smith, paleontologist at University of Wisconsin
“The presence of uniramous limbs in an aquatic ancestor suggests these features were not solely adaptations for terrestrial life but may have been pre-existing traits that facilitated the eventual land colonization.”
— Lead researcher Dr. Robert Egan

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What Remains Unclear
It remains unclear exactly how widespread such aquatic ancestors were among early myriapods and whether similar limb traits existed in other lineages. The precise ecological role of the head appendages and the full range of behaviors of Waukartus are still unknown. Further fossil discoveries are needed to clarify these aspects.

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What’s Next
Future research will focus on finding more fossils of similar age and morphology to better understand the diversity of early aquatic myriapod ancestors. Additional analyses may clarify how limb reduction and habitat shifts co-evolved, and whether other lineages experienced similar pre-adaptations.
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Key Questions
What does this discovery tell us about the evolution of millipedes and centipedes?
It shows that their ancestors already had limb structures suited for aquatic environments before moving onto land, indicating a more complex evolutionary process involving pre-adaptations.
How does this change previous ideas about limb evolution in myriapods?
It challenges the view that limb reduction was solely a terrestrial adaptation, suggesting it occurred earlier in aquatic ancestors and was an exaptation for land life.
Are there other fossils like Waukartus muscularis?
So far, Waukartus is a unique find, but ongoing excavations may uncover more specimens that shed light on early myriapod evolution.
What is an exaptation?
An exaptation is an evolutionary trait that originally evolved for one purpose but was later co-opted for a different function, such as limb reduction in aquatic ancestors aiding terrestrial adaptation.