Monotremes as Pretribosphenic Mammals

An abundance of evidence points to the conclusion that monotremes are phyletically allied with pretribosphenic, rather than with tribosphenic, mammals. Monotremes do not have a tribosphenic dentition. Character analyses that apply tribosphenic cusp terminology to monotreme dentitions are implicitly limited thereby. A review of the molar dentition of living and fossil monotremes suggests that upper molars are composed of a strongly developed pretribosphenic paracone and metacone and a series of stylar cusps attached to them in a bicrescentic, or dilambdodont, fashion. The lower molars are composed of a trigonid, with a pretribosphenic protoconid, paraconid, and metaconid, and distal metacristid. The paraconid of m1 is reduced or lost. The talonid is composed of the pretribosphenic hypoconid, hypoconulid, and cristid obliqua. There is no evidence for a tribosphenic entoconid, nor for a talonid basin. There was no tribosphenic protocone. Monotremes are not related to other taxa included in Australosphenida. The dentition of Cretaceous taxa, such as Teinolophos and Steropodon, apparently still functioned by orthal mechanisms, whereas by the medial Paleocene (Monotrematum) and later (Obdurodon), monotremes appear to have accommodated a diet of soft-bodied organisms that left little trace of a mastication regime that had changed to apical wear via propalinal motion. Monotremes appear to be modern representatives of a Mesozoic radiation of pretribosphenic mammals centered largely in Gondwana, where they still reside today.     

A fossil baboon skull from the lower Omo basin,southwest Ethiopia

A well-preserved fossil skull assignable to the genusPapio was discovered in the late Pliocene deposits at the “White Sands locality” of the lower Omo basin, southwest Ethiopia in November 1978. This skull obviously belongs to a male baboon which is morphologically somewhat different from the extant species ofPapio and closely resemblesPapio baringensis R. E. F. Leakey, 1969. Its muzzle is not only long but comparatively wide and robust with a very flat dorsum, an almost quadrate vertical cross section, short and obtuse maxillary ridges and virtually absent maxillary fossae, and its zygomatic portion flares laterally to widen the large temporal process of the zygomatic bone. The post-orbital constriction is very marked, the temporal fossa is squarish in basal view, and the sagittal crest is placed rather posteriorly and meets a large nuchal crest, all of which suggest heavy temporal musculature. The incisors are lacking but were probably not as small as inP. baringensis. As a result of morphological comparisons with extant and fossil species ofPapio, this newly discovered skull was considered to represent a new species,Papio quadratirostris.     

Late Jurassic Mammals from Tendaguru, Tanzania, East Africa

Records of Mesozoic mammals are extremely rare in Africa. The only previous record from the Upper Jurassic of Africa is a fragmentary mandible without teeth of Brancatherulum tendagurense. Here I report the discovery of two new mammals from the Upper Jurassic of Tendaguru, Tanzania. The fossils were recovered from the Middle Saurian Bed of the Tendaguru Series. A lower molar of a triconodontid mammal is described as Tendagurodon janenschi gen. et sp. nov., and a fragmentary dentary of a eupantothere as Tendagurutherium dietrichi gen. et sp. nov. The eupantothere in particular contributes to documenting the evolution of mammals during the Mesozoic. The posterior portion of the mandible of Tendagurutherium dietrichi gen. et sp. nov. shows that the angular (tympanic) bone was not yet completely separated from the dentary, a previously undocumented stage of eupantotherian middle ear evolution.     

A new taeniolabidoid multituberculate (Mammalia) from the middle Puercan of the Nacimiento Formation,New Mexico,and a revision of taeniolabidoid systematics and phylogeny

Multituberculates were amongst the most abundant and taxonomically diverse mammals of the late Mesozoic and the Palaeocene, reaching their zenith in diversity and body size in the Palaeocene. Taeniolabidoidea, the topic of this paper, includes the largest known multituberculates, which possess highly complex cheek teeth adapted for herbivory. A new specimen from the early Palaeocene (middle Puercan; biochron Pu2) of the Nacimiento Formation, New Mexico represents a new large‐bodied taeniolabidoid genus and species, K imbetopsalis simmonsae . A phylogenetic analysis to examine the relationships within Taeniolabidoidea that includes new information from K imbetopsalis gen. et sp. nov. and gen. nov. and from new specimens of Catopsalis fissidens, first described here, and data from all other described North American and Asian taeniolabidoids. This analysis indicates that Catopsalis is nonmonophyletic and justifies our transfer of the basal‐most taeniolabidoid ‘Catopsalis’ joyneri to a new genus, V alenopsalis . K imbetopsalis and Taeniolabis form a clade (Taeniolabididae), as do the Asian Lambdopsalis, Sphenopsalis, and possibly also Prionessus (Lambdopsalidae). Taeniolabidoids underwent a modest taxonomic radiation during the early Palaeocene of North America and underwent a dramatic increase in body size, with Taeniolabis taoensis possibly exceeding 100 kg. Taeniolabidoids appear to have gone extinct in North America by the late Palaeocene but the appearance of lambdopsalids in the late Palaeocene of Asia suggests that they dispersed from North America in the early to middle Palaeocene.     

The cephalic morphology of the troglobiontic cholevine species Troglocharinus ferreri (Coleoptera,Leiodidae)

Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species Troglocharinus ferreri (Cholevinae, Leptodirini) was examined in detail. Overall, the general pattern is similar to what is found in Catops ventricosus (Cholevini). Shared apomorphic features include a fully exposed anterolateral concavity containing the antennal socket, a distinct bead above this depression, a bilobed lip-like structure anterad the labrum, a flat elevated portion of the ventral mandibular surface, and a ventral process at the proximomesal edge of this mandibular area. The tentorial structures are well-developed as in C. ventricosus, with a large laminatentorium and somewhat shortened dorsal arms. The mouthparts are largely unmodified, with the exception of unusually well-developed extrinsic maxillary muscles. Features of T. ferreri obviously linked with subterranean habits are the complete lack of compound eyes, circumocular ridges, and optic lobes. A series of characters is similar to conditions found in other genera of Leptodirini: the head capsule completely lacks a protruding ocular region, a distinct neck is missing, the transverse occipital crest is indistinct, and the antennae are elongate and lack a distinct club. Two different trends of cephalic transformations occur in troglobiontic Leptodirini, with some genera like Troglocharinus and Leptodirus having elongated head capsules and antennae, and others having broadened, more transverse heads. In contrast, the modifications are more uniform in the closely related Ptomaphagini, with a pattern distinctly differing from Leptodirini: the head is transverse, with a distinctly protruding ocular region, a distinct transverse occipital crest, and a very narrow neck region.     

New Studies of Enamel Microstructure in Mesozoic Mammals: A Review of Enamel Prisms as a Mammalian Synapomorphy

Characters from enamel microstructure have not been used in recent phylogenetic analyses of Mesozoic Mammalia. Reasons are that enamel characters have been perceived as (A) variable without regard to systematic position of taxa, (B) inconsistently reported within the literature, and (C) simply scored as either prismatic or not prismatic in earlier mammals. Our work on Mesozoic mammals such as Sinoconodon, Gobiconodon, Triconodontidae, Docodon, Laolestes, and others suggests that synapsid columnar enamel (SCE) structure was easily transformed into plesiomorphic prismatic enamel (PPE) and that PPE may be described with at least five independent character states. Two PPE characters—a flat, open prism sheath and a planar prism seam—were present in the cynodont Pachygenelus and in several Jurassic and Cretaceous mammals. We propose that appearance of a prism sheath transforms SCE into PPE and that reduction and loss of a prism sheath reverse PPE into SCE, in both phylogeny and ontogeny. We further propose that no amniote vertebrates other than the trithelodontid cynodont, Pachygenelus, plus Mammalia have ever evolved an ameloblastic Tomes process capable of secreting PPE and that the genetic potential to secrete PPE is a synapomorphy of Pachygenelus plus Mammalia, whether or not all lineages of the clade have expressed that potential.     

A unique algal ridge system in the Exuma Cays, Bahamas

 An algal ridge system discovered along the Exuma Cays, Bahamas constructs bioherms to a thickness of at least 1.5 m and is associated with modern intertidal stromatolites. These algal ridges are unique because they grow in atypical environments characterized by relatively low wave energy, high rates of sedimentation and low rates of herbivory. They also are composed primarily of the branching crustose coralline alga, Neogoniolithon strictum, which heretofore was not known to form algal ridges. Lateral growth rates of crusts, vertical growth rates of branches and survivorship of transplanted N. strictum were greatest in the shallow fore reef zone of the algal ridge. The alga is also capable of surviving and growing when covered with sediment for at least 100 days. Under such conditions it transforms from a branched to an unbranched morphology. Parrotfish grazing, which is said to limit the abundance of branched corallines and algal ridges, was two orders of magnitude lower than in published accounts from other reef systems of the Caribbean and one order of magnitude less than that found on nearby coral reefs of the west Exuma Sound. Neogoniolithon strictum, a delicate and open-branched coralline, persisted for over a year without grazing damage when transplanted to a depth of 2.3 m. This algal ridge-building coralline becomes a well-indurated limestone following submarine lithification of sediment that infills the open branch framework. As a result, N. strictum ridges are comparable to the dense frameworks associated with most algal ridges. Observations of N. strictum -associated bioherms along Central America suggest this ridge system may exist elsewhere under conditions similar to those described for the Bahamas. Accepted: 23 March 1996     

Revised paleoenvironmental analysis of the Holocene portion of the Barbados sea-level record: Cobbler’s Reef revisited

Sample elevations corrected for tectonic uplift and assessed relative to local modeled sea levels provide a new perspective on paleoenvironmental history at Cobbler’s Reef, Barbados. Previously, ¹⁴C-dated surface samples of fragmented Acropora palmata plotted above paleo sea level based on their present (uplifted) elevations, suggesting supratidal rubble deposited during a period of extreme storms (4500–3000 cal BP), precipitating reef demise. At several sites, however, A. palmata persisted, existing until ~370 cal BP. Uplift-corrected A. palmata sample elevations lie below the western Atlantic sea-level curve, and ~2 m below ICE-6G-modeled paleo sea level, under slow rates of sea-level rise, negating the possibility that Cobbler’s Reef is a supratidal storm ridge. Most sites show limited age ranges from corals likely damaged/killed on the reef crest, not the mixed ages of rubble ridges, strongly suggesting the reef framework died off in stages over 6500 yr. Reef crest death assemblages invoke multiple paleohistoric causes, from ubiquitous hurricanes to anthropogenic impacts. Comparison of death assemblage ages to dated regional paleotempestological sequences, proxy-based paleotemperatures, recorded hurricanes, tsunamis, European settlement, deforestation, and resulting turbidity, reveals many possible factors inimical to the survival of A. palmata along Cobbler’s Reef.

     

Two new species of the Trichostrongylidae (Nematoda: Trichostrongyloidea), parasitic in Gorilla gorilla beringei in Uganda

Two new species of trichostrongylid parasites of a mountain gorilla in Uganda are described. Paralibyostrongylus kalinae n. sp. is characterised by a short dorsal ray, a dorsal oesophageal tooth and two median longitudinal ridges in the vulvar region. The species is more closely related to leporid parasites than to P. hebrenicutus, another species known from the gorilla. A “transfuge” phenomenon, i.e. transfer to a new host without speciation is suggested. Hyostrongylus kigeziensis n. sp. has several specific morphological characteristics and in particular a synlophe with many strong ridges, rays 4 and 5 divergent at the apex, and the dorsal ray with two branches. Hyostrongylus appears to be an Ethiopian genus which has radiated in many archaic mammals. With the technical collaboration of Mrs. N. Caillaud and Mr R. Tchesprakoff. With the technical collaboration of Mrs. N. Caillaud and Mr R. Tchesprakoff.     

SYSTEMATIC REVISION OF PSEUDOBEACONIA BORDAS, 1944, AND MENDOCINICHTHYS WHITLEY, 1953 (ACTINOPTERYGII: ‘PERLEIDIFORMES’) FROM THE TRIASSIC OF ARGENTINA

Abstract: The ‘perleidiform’Mendocinichthys and Pseudobeaconia from the Potrerillos and Santa Clara Abajo formations (Upper Triassic; Argentina) are reviewed. Mendocinichthys has been known from a review of this species that is not based on the type material, but on referred specimens in the American Museum of Natural History. However, those specimens are found here to represent a new species of Pseudobeaconia, P. celestae sp. nov. Consequently, Mendocinichthys is restricted here to the type material and, within it, to the only specimen that clearly represents a distinct taxon and is thus designated lectotype. We further performed a cladistic analysis and propose a new family, Pseudobeaconiidae for these two South American Triassic genera, which is mainly characterized by the presence of an incomplete dorsal ridge of spine‐like scales between the skull and the dorsal fin, and scales with straight posterior border, an elevated central region and marginal concentric ridges of ganoine. The cladistic analysis further indicates the existence of some lineages endemic to certain areas of Gondwana and Europe. Pseudobeaconia celestae sp. nov. represents the first record of Pseudobeaconia in the Cacheuta sub‐basin of the Cuyana Basin. The genus was previously known from the Santa Clara sub‐basin of the Cuyana Basin only, and the new record confirms the previous hypothesis of correlation between the sedimentary infilling of these sub‐basins.     

Implications of rudd (Scardinius erythrophthalmus) herbivory on submerged macrophytes in a shallow eutrophic lake

The role of rudd (Scardinius erythrophthalmus) herbivory was studied in Kirkkojärvi, a shallow and turbid basin in Lake Hiidenvesi, Finland. The submerged species dominating in the rudd diets were Potamogeton obtusifolius, Ranunculus circinatus, Sparganium emersum, bryophytes, and filamentous algae. Plant consumption estimated with bioenergetics modelling increased with fish age, being highest in late summer concomitant with the macrophyte biomass peak. Depending on the age structure, a rudd biomass of 20 kg ha^?1 consumes 18–23 kg of macrophytes ha^?1 a^?1, while a rudd biomass of 100 kg ha^?1 results in plant consumption of 92–115 kg ha^?1 a^?1. Although, rudd seemed to feed rather unselectively on suitable-sized and edible plants, some species abundant in the littoral, such as Myriophyllum verticillatum and pleustophytic Ceratophyllum demersum, were not found in rudd guts, indicating selective plant consumption. In Kirkkojärvi, selective grazing by rudd and increased turbidity and high nutrient levels partly caused by bottom dwelling cyprinid fish, may promote the inedible and pleustophytic macrophytes, which have increased in Kirkkojärvi during the past decades.

     

New ichnospecies of Linichnus Jacobsen & Bromley, 2009

Abstract

A new ichnospecies, Linichnus bromleyi, is described on bone substrate as the result of a very likely predator/scavenger interaction. L. bromleyi consists of a single groove with a non-serrated edge. This new ichnotaxon is compared with L. serratus which was defined as a single elongate serrate-edged groove. L. bromleyi has been identified over the surface of bones of marine mammals (in particular, cetaceans and pinnipeds) from two Pliocene outcrops of Andalusia, southern Spain. The new ichnospecies can be clearly related with a trophic interaction between sharks and marine mammals (mainly whales, dolphins and seals).     

A new species of Dermopristis Kearn, Whittington & Evans-Gowing, 2010 (Monogenea: Microbothriidae), with observations on associations between the gut diverticula and reproductive system and on the presence of denticles in the nasal fossae of the host Glaucostegus typus (Bennett) (Elasmobranchii: Rhinobatidae)

Dermopristis cairae n. sp. (Monogenea: Microbothriidae) is described from the skin and possibly from the nasal fossae of the giant shovel-nosed ray Glaucostegus typus (Bennett). The new species is distinguished from D. paradoxus Kearn, Whittington & Evans-Gowing, 2010 by its larger size, body shape, lack of transverse ridges on the ventral surface and absence of a seminal receptacle. Extensive short gut branches lie dorsal to the testes and adjacent to the coiled region of the vas deferens and the o?type, possibly reflecting high metabolic demand in these areas. Denticles are present in the lining of the nasal fossae of G. typus, providing a firm substrate for the cement-based attachment of a microbothriid. However, confirmation that D. cairae inhabits the nasal fossae of G. typus is required.     

The morphology and wear of the lingual notch in macaques and langurs

Although interspecific differences in the relative development of molar occlusal features have been documented for most primate species, wear-related changes in these features are only beginning to be understood. In the present study, the notch between the metaconid and entoconid of M₂ was examined in a skeletal sample of Macaca fascicularis, Presbytis cristatus, and Presbytis rubicunda from the Museum of Comparative Zoology at Harvard University. Photogrammetric analyses yielded X, Y, and Z coordinates that were used to compute three angular measurements and two length measurements at the lingual notch. Statistical analyses of measurements from unworn dentitions indicated that the mesial slope of the entoconid is steeper than the distal slope of the metaconid in all three species. In addition, interspecific comparisons demonstrated that P. cristatus has the steepest lingual notch bordered by the largest molar shearing crests, while P. rubicunda and M. fascicularis have molar shearing crests of similar size, with P. rubicunda merely having higher cusps that are closer together than those of M. fascicularis. Fanally, P. cristatus shows a smaller decrease in lingual cusp relief with wear than M. fascicularis, while P. cristatus and P. rubicunda probably show similar decreases in lingual cusp relief with wear. The differences in wear-related changes in lingual cusp relief between M. fascicularis and P. cristatus may well be related to the frugivorous/folivorous dietary differences between these species.     

History of exposure to herbivores increases the compensatory ability of an invasive plant

Release from natural enemies is frequently cited as an important factor contributing to plant invasions. But such effects are likely to be temporary—native herbivores can form new plant-herbivore associations and co-evolved insects might reach the new range. While the potential effects of the initial enemy release have been well studied, the consequences of any resumption of herbivory are poorly understood. Alternanthera philoxeroides is one of the most widespread invasive plants in China and is attacked both by a specialist herbivore introduced from the native range, Agasicles hygrophila, and a native beetle Cassida piperata Hope which has formed a new association. However, these insects are not found throughout the invaded range. To test the effect of the history of population exposure to herbivory on compensatory ability, plants were cultured from 14 populations around China that differed in whether A. hygrophila or C. piperata were present. Treatment plants were exposed to herbivory by A. hygrophila for a week until 50% of the leaf area was defoliated, then grown for 80 days. Plants from populations with prior exposure to herbivory (of any kind) accumulated more root mass than populations without prior exposure, indicating that prior exposure to insects can stimulate plant compensation to herbivory. We would recommend that potential changes in plant tolerance in response to prior exposure to herbivory are considered in invasive plant management plans that employ bio-control agents.     

Contribution of flexible allocation priorities to herbivory tolerance in C4 perennial grasses: an evaluation with 13C labeling

The ability of plants to rapidly replace photosynthetic tissues following defoliation represents a resistance strategy referred to as herbivory tolerance. Rapid reprioritization of carbon allocation to regrowing shoots at the expense of roots following defoliation is a widely documented tolerance mechanism. An experiment was conducted in a controlled environment to test the hypothesis that herbivory-sensitive perennial grasses display less flexibility in reprioritizing carbon allocation in response to defoliation than do grasses possessing greater herbivory tolerance. An equivalent proportion of shoot biomass (60% dry weight) was removed from two C₄ perennial grasses recognized as herbivory-sensitive, Andropogon gerardii and Schizachyrium scoparium, and two C₄ perennial grasses recognized as herbivory-tolerant, Aristida purpurea and Bouteloua rigidiseta. Both defoliated and undefoliated plants were exposed to ¹³CO₂ for 30 min, five plants per species were harvested at 6, 72 and 168 h following labeling, and biomass was analyzed by isotope ratio mass spectrometry. The tallgrass, A. geraiddii, exhibited inflexible allocation priorities while the shortgrass, B. rigidiseta, exhibited flexible allocation priorities in response to defoliation which corresponded with their initial designations as herbivory-sensitive and herbivory-tolerant species, respectively. A. gerardii had the greatest percentage and concentration of ¹³C within roots and lowest percentage of ¹³C within regrowth of the four species evaluated. In contrast, B. rigidiseta had a greater percentage of ¹³C within regrowth than did A. gerardii, the greatest percentage of ¹³C within new leaves of defoliated plants, and the lowest concentration of ¹³C within roots follwing defoliation. Although both midgrasses, S. scoparium and A. purpurea, demonstrated flexible allocation priorities in response to defoliation, they were counter to those stated in the initial hypothesis. The concentration of ¹³C within new leaves of S. scoparium increased in response to a single defoliation while the percentage and concentration of ¹³C within roots was reduced. A. purpurea was the only species in which the percentate of ¹³C within new leaves decreased while the percentage of ¹³C within roots increased following defoliation. The most plausible alternative hypothesis to explain the inconsistency between the demonstrated responsiveness of allocation priorities to defoliation and the recognized herbivory resistance of S. scoparium and A. purpurea is that the relative ability of these species to avoid herbivory may make an equal or greater contribution to their overall herbivory resistance than does herbivory tolerance. Selective herbivory may contribute to S. scoparium's designation as a herbivorysensitive species even though it possesses flexible allocation priorities in response to defoliation. Alternatively, the recognized herbivory resistance of A. purpurea may be a consequence of infrequent and/or lenient herbivory associated with the expression of avoidance mechanisms, rather than the expression of tolerance mechanisms. A greater understanding of the relative contribution of tolerance and avoidance strategies of herbivory resistance are required to accurately interpret how herbivory influences plant function, competitive interactions, and species abundance in grazed communities.     

A new blind loach, <Emphasis Type="Italic">Oreonectes elongatus</Emphasis> sp. nov. (Cypriniformes: Balitoridae) from Guangxi,China

A new loach, Oreonectes elongatus sp. nov. is described based on collections from Mulun Township, Huanjiang County, Guangxi in China. It is distinguished from its congeners by the combination of the following characters: most elongate body (body depth/SL 8.62–10.68%), blind, a forked caudal fin, obvious adipose dorsal crest and ventral crest; entire body naked and de-pigmented. Although the new species has a similar distribution with O. macrolepis, it can be distinguished by scales (absent in O. elongatus vs. present in O. macrolepis), shape of snout (elongate vs. round), the opposite position of the dorsal and pelvic fins origins (behind vs. front). The new species shares the same possession of dorsal and ventral crests, a forked caudal fin, eyeless, naked body and incomplete lateral line with O. translucens, but can be distinguished from the latter by caudal fin crest (more developed and translucent in O. translucens), longer anterior nostril tube and barbel, extreme of pectoral fin reaching 2/3 of the distance between origin of pectoral and pelvic fins, more vertebrae (4 + 38–39 vs. 4 + 32).     

Early Cretaceous non‐marine Ostracoda from the North Falkland Basin,South Atlantic

Twenty‐seven species of well‐preserved and abundant Early Cretaceous non‐marine ostracod crustaceans were recovered from the North Falkland Basin. The assemblage is unusually diverse for a non‐marine palaeoenvironment and is sourced from cuttings samples collected during 2011 drilling of wells by Desire and Rockhopper Exploration, in the northern and southern areas of the basin. Ostracoda are entirely undocumented in published accounts from this basin, and all but one species appear to be new to science. For the new taxa, one new family (Alloiocyprideidae; type genus Hourcqia) is proposed and includes Hourcqia woodi sp. nov. Four new genera are erected: Falklandicypris gen. nov.; type species F. petrasaltata sp. nov., Gangamoncythere gen. nov.; type species G. colini sp. nov., Paraplesiocypridea gen. nov.; type species P. alloios sp. nov., and Musacchiocythere gen nov.; type species M. sarunata (Musacchio, 1978 ). Nine other species are described as new and are Falklandicypris desiderata, Clinocypris epacrus, Cypria poietes, Ilhasina? leiodermatus, Looneyellopsis tuberculatus, Theriosynoecum petasmathylacus, Theriosynoecum ballentae, Timiriasevia fluitans, and Vecticypris samesi. The remainder are left in open nomenclature due to paucity of material. The ostracod assemblage is largely restricted to the southern part of the basin (wells 25/5‐1 and 26/6‐1). In the northern part of the basin only four species, dominated by Vecticypris samesi, are present and with one possible exception are restricted there. There appears to have been little or no interchange of species suggesting that a barrier probably existed between the northern and southern regions. The more diverse southern assemblage indicates that more favourable conditions existed to the south. Stratigraphically, a distinct change in faunal composition recorded in both southern wells is likely to be an isochronous event correlatable across the southern area, and of an age no younger than Hauterivian.     

Untangling individual variation in natural populations: ecological, genetic and epigenetic correlates of long-term inequality in herbivory

Individual variation in ecologically important features of organisms is a crucial element in ecology and evolution, yet disentangling its underlying causes is difficult in natural populations. We applied a genomic scan approach using amplified fragment length polymorphism (AFLP) markers to quantify the genetic basis of long‐term individual differences in herbivory by mammals at a wild population of the violet Viola cazorlensis monitored for two decades. In addition, methylation‐sensitive amplified polymorphism (MSAP) analyses were used to investigate the association between browsing damage and epigenetic characteristics of individuals, an aspect that has been not previously explored for any wild plant. Structural equation modelling was used to identify likely causal structures linking genotypes, epigenotypes and herbivory. Individuals of V. cazorlensis differed widely in the incidence of browsing mammals over the 20‐year study period. Six AFLP markers (1.6% of total) were significantly related to herbivory, accounting altogether for 44% of population‐wide variance in herbivory levels. MSAP analyses revealed considerable epigenetic variation among individuals, and differential browsing damage was significantly related to variation in multilocus epigenotypes. In addition, variation across plants in epigenetic characteristics was related to variation in several herbivory‐related AFLP markers. Statistical comparison of alternative causal models suggested that individual differences in herbivory are the outcome of a complex causal structure where genotypes and epigenotypes are interconnected and have direct and indirect effects on herbivory. Insofar as methylation states of MSAP markers influential on herbivory are transgenerationally heritable, herbivore‐driven evolutionary changes at the study population will involve correlated changes in genotypic and epigenotypic distributions.