Habeliida is an order of extinct arthropods that existed during the middle Cambrian. They are thought to be some of the oldest members of the chelicerate lineage.

Description

Members of Habeliida are characterized by a numbers of features: These include: bulging head shields, which have notches to accommodate a pair of compound eyes, and triangular extensions outward to the sides. Attached to the head are 5 pairs of segmented, thin upper antennae-like exopods (upper limb branches), with along with a reduced first appendage at least in Habelia, followed by 5 pairs of seven-segmented spined endopod (lower branch) grasping appendages, which are attached to basal segments (basipods), that increase in size posteriorly and are gnathobasic, that is modified to have inward-facing spines. Along the body run limbs with paddle-shaped exopods which are fringed by narrow lamellae. The body ends with a telson, which is either elongate and spine like (Habelia) or paddle-shaped (Sanctacarididae).

Ecology

Restoration of Wisangocaris feeding on trilobites, as inferred from stomach contents

Habeliids were likely predatory, using the upper exopod head appendages as sensory organs, while using the lower spined head endopods to grasp prey. They are thought to have been durophagous, with the robust gnathobases (spined basal sections of limbs) used to shred hard-shelled organisms. Remains of trilobites have been found as stomach contents in Wisangocaris.

Taxonomy

It is divided into two families, Habeliidae (monotypic, containing only Habelia), and Sanctacarididae (containing Sanctacaris, Utahcaris and Wisangocaris) Messorocaris has been suggested to be part of the order in some studies, but this is uncertain. They are suggested to be stem-group chelicerates, though they lack the chelicerae present in true chelicerates. They may be the ancestral group from which modern chelicerates emerged:

Cladogram after O’Flynn et al, 2023:

Total group Arthropoda"Gilled lobopodians" (Pambdelurion, Kerygmachela) Opabinia Radiodonta (e.g Anomalocaris) Deuteropoda Kylinxia Fengzhengia Kiisortoqia Bushizheia Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) "Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
"Gilled lobopodians" (Pambdelurion, Kerygmachela)
Opabinia Radiodonta (e.g Anomalocaris) Deuteropoda Kylinxia Fengzhengia Kiisortoqia Bushizheia Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) "Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Opabinia
Radiodonta (e.g Anomalocaris) Deuteropoda Kylinxia Fengzhengia Kiisortoqia Bushizheia Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) "Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Radiodonta (e.g Anomalocaris)
DeuteropodaKylinxia Fengzhengia Kiisortoqia Bushizheia Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) "Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Kylinxia Fengzhengia
Kylinxia
Fengzhengia
Kiisortoqia Bushizheia Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) "Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Kiisortoqia Bushizheia
Kiisortoqia
Bushizheia
Megacheira Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc)
Megacheira
Habeliida Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc)
Habeliida
Mollisonia Chelicerata (horseshoe crabs, sea spiders, arachnids, etc)
Mollisonia
Chelicerata (horseshoe crabs, sea spiders, arachnids, etc)
"Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
"Great appendage bivalved forms" (Occacaris, Forfexicaris) Isoxyida
"Great appendage bivalved forms" (Occacaris, Forfexicaris)
Isoxyida
Artiopoda (including Trilobita) Mandibulata Fuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Artiopoda (including Trilobita)
MandibulataFuxianhuiida Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Fuxianhuiida
Myriapoda (millipedes, centipedes, etc) Hymenocarina Pancrustacea (crustaceans, insects, etc)
Myriapoda (millipedes, centipedes, etc)
Hymenocarina Pancrustacea (crustaceans, insects, etc)
Hymenocarina
Pancrustacea (crustaceans, insects, etc)

Cladogram after Lerosey-Aubril and Ortega-Hernández (2026):

Pan‑Chelicerata †JianfengiaSklerolibyon MegacheiraFortiforcepsYohoiaHaikoucarisLeanchoiliidaeMollisoniidaWisangocaris Habeliida †HabeliaSanctacarisUtahcaris chelicerae present †MegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Pan‑ChelicerataJianfengiaSklerolibyon MegacheiraFortiforcepsYohoiaHaikoucarisLeanchoiliidaeMollisoniidaWisangocaris Habeliida †HabeliaSanctacarisUtahcaris chelicerae present †MegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
JianfengiaSklerolibyonMegacheira
Jianfengia
Sklerolibyon
FortiforcepsYohoiaHaikoucarisLeanchoiliidaeMollisoniidaWisangocaris Habeliida †HabeliaSanctacarisUtahcaris chelicerae present †MegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Fortiforceps
Yohoia
HaikoucarisLeanchoiliidae
Haikoucaris
Leanchoiliidae
MollisoniidaWisangocaris Habeliida †HabeliaSanctacarisUtahcaris chelicerae present †MegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Mollisoniida
Wisangocaris Habeliida †HabeliaSanctacarisUtahcaris chelicerae present †MegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
WisangocarisHabeliida
Habelia
SanctacarisUtahcaris
Sanctacaris
Utahcaris
chelicerae presentMegacheliceraxOffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Megachelicerax
OffacolusSetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Offacolus
SetapeditesDibasteriumWeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
SetapeditesDibasterium
Setapedites
Dibasterium
WeinberginaVenustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Weinbergina
Venustulus Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Venustulus
Pycnogonida (sea spiders) Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Pycnogonida (sea spiders)
Xiphosura (horseshoe crabs) ArachnidaEurypterida (sea scorpions)
Xiphosura (horseshoe crabs)
ArachnidaEurypterida (sea scorpions)
Arachnida
Eurypterida (sea scorpions)