Horseshoe Crab: the living fossil

Introduction

Witness to almost 500 million years of Earth's history, horseshoe crabs are the textbook definition of living fossils. Their alien-like primitive forms, featuring the iconic spine-like telson and horseshoe-shaped carapace, have remained virtually unchanged across their existence. While their resemblance to a horseshoe is difficult to challenge, “crab” is a misnomer for the creatures considering they are not even crustaceans. They are, in fact, more closely related to spiders and scorpions.

Present-day horseshoe crabs are all marine dwellers although forays into freshwater have been noted among long extinct groups. The mangrove horseshoe crab, at odds with the other three extant species, also inhabits the brackish waters found near mangrove forests.

Figure 1. Front and back views of Limulus polyphemus showing general morphology

Phylogeny and Evolution

Modern-day horseshoe crabs are chelicerate arthropods of the Limulidae family. Despite their name, they are not crustaceans at all and are, in fact, more closely related to arachnids. Of 88 formally described species, only 4 remain today and are the only surviving members of the order Xiphosura. The extant species include the Atlantic horseshoe crab Limulus polyphemus, and three Asiatic species: the Indo-Pacific horseshoe crab Tachypleus gigas, the tri-spine horseshoe crab Tachypleus tridentatus, and the mangrove horseshoe crab Carcinoscorpius rotundicauda.

The first Xiphosurans emerged some 470 million years ago in the Lower Ordovician period of the Paleozoic era, making horseshoe crabs one of the most ancestral species of animals still alive today. The Limulids, the family of modern horseshoe crabs, first appeared around 250 years ago in the Triassic period. Two rounds of evolutionary stasis have been identified in the Xiphosurans: the conservation of prosomal and opisthosomal growth rates near their origin in the Lower Ordivician and the conservation of shape in the Jurassic period owing to the emergence of the Limulids in the Triassic. Notwithstanding physiological changes, present-day crabs are larger in size than their ancestral counterparts and have their abdominal segments fused together.

Morphology

All horseshoe crabs share the same body plan: the cephalothorax or prosoma (head and chest fused together), the opisthosoma (the abdomen with the inclusion of the heart and respiratory organs, a distinctive feature of the chelicerates) and, the iconic feature of the horseshoe crab, the telson (the long tail-like spine jutting out from the abdomen). The prosoma is shrouded in a protective carapace bearing an uncanny resemblance to a horseshoe, giving the horseshoe crab its name. Unlike any other extant chelicerate, horseshoe crabs possess, among its ten eyes, two compound eyes that sit atop its carapace; they also have the largest rod and cone cells of any known animal.

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Edited by [Author Name], student of Joan Slonczewski for BIOL 116, 2024, Kenyon College.