Nematodes are one of the most abundant groups of invertebrates on the face of the earth. Their extremely poor fossil record hinders our ability to assess just when members of this group invaded land and first became associated with plants. This study reports fossil nematodes from the stomatal chambers of the Early Devonian (396 mya) land plant, Aglaophyton major. These nematodes, which are tentatively assigned to the order Enoplia, are described as Palaeonema phyticum gen. n., sp. n. in the new family Palaeonematidae fam. n. Diagnostic characters of the family are: i) cuticular striations; ii) uniform, cylindrical pharynx with the terminal portion only slightly set off from the remainder; and iii) a two-portioned buccal cavity with the upper portion bearing protuberances. The presence of eggs, juveniles and adults in family clusters within the plant tissues provide the earliest evidence of an association between terrestrial plants and animals and may represent an early stage in the evolution of plant parasitism by nematodes.