The growth of knowledge and research practices in any discipline is characterised by a trade-off between depth and breadth: we can either invest efforts to learn a little about many things, or learn a lot about few things. In parasitology, breadth of knowledge corresponds to research on biodiversity and taxonomy: the discovery and description of an increasing number of new species. In contrast, depth of knowledge comes from focused research on a few model species, about which we accumulate much detailed information. Breadth and depth of knowledge are equally important for progress in parasitology. In this essay, focusing on trematodes, I demonstrate that current research is rapidly broadening our knowledge (high rate of new trematode species being discovered) but not deepening that knowledge at a comparable rate. The use of model species, with caveats, appears to offer a promising avenue for deeper knowledge. I present a case study illustrating how it is possible to develop new model trematode species at low cost to increase the depth of our understanding in areas including host-parasite ecological dynamics, co-evolution, and responses to environmental and climatic changes. The take-home message serves as a call to action to parasitologists, emphasising the need to focus as much effort on depth of knowledge as we currently invest in breadth of knowledge.

New, well-known and predicted life cycles for trematodes of the Haploporoidea (Haploporidae and Emprostiotrematidae) and three families of the Lepocreadioidea (Enenteridae, Gorgocephalidae, Gyliauchenidae) involve encystment of the metacercaria in the open (usually on vegetation) followed by ingestion by a range of herbivorous or detritivorous fishes. These life cycles appear among relatively highly derived plagiorchiidan trematodes in which three-host life cycles incorporating an animal second intermediate host are dominant. We hypothesise that the two-host life cycles in the Haploporoidea and Lepocreadioidea arose by secondary truncation of a three-host cycle; the second intermediate host was lost in favour of encystment in the open. Modification of a three-host life cycle effective for the infection of carnivores is consistent with the understanding that fishes arose as carnivores and that multiple lineages have secondarily become detritivores and herbivores. Four of the five trematode families involved infect fishes relating to multiple orders, suggesting a complex history of host-switching. In contrast, the Gorgocephalidae, the smallest of the families, has been found only in a single family, Kyphosidae. The timing of the evolutionary events leading to this putative life cycle truncation is yet to be deduced, but the rich developing understanding of the history of the fishes creates a strong template for future analysis.

Iceland is an isolated, sub-Arctic, oceanic island of volcanic origin in the northern North Atlantic. With a limited faunal diversity and being the most northern point in the distributional range for some species, it is an intriguing model region to study parasite biodiversity and biogeography. Since 2006, there has been a history of intense biodiversity discoveries of freshwater trematodes (Trematoda, Digenea), thanks to the use of integrative taxonomic methods. The majority of digeneans (28 out of 41 known) were characterised with molecular genetic methods and morphological analyses, with some of their life-cycle stages and geographical distribution assessed. A surprising diversity has been discovered, comprising species of the families Allocreadiidae, Cyclocoeliidae, Diplostomidae, Echinostomatidae, Gorgoderidae, Plagiorchiidae, Notocotylidae, Schistosomatidae, and Strigeidae. Many of the recorded species complete their life cycles within Iceland, with three snail species (Ampullaceana balthica, Gyraulus parvus, Physa acuta) known as intermediate hosts. No trematodes endemic for Iceland were found; they appear to be generalists with wide geographical ranges dispersed mainly by migratory birds. Interestingly, fish trematodes recorded in Iceland were found in mainland Europe, indicating that they might be dispersed by anadromous fishes, by human activity, or by migratory birds carrying intermediate hosts. The trematode fauna is mainly Palaearctic, with few species recorded in North America. We highlight the ongoing need for precise species identification via integrative taxonomic methods, which is a baseline for any further ecological studies and adequate epidemiological and conservation measures. Also, there is still a need of obtaining well-preserved vouchers of adults for definite species delimitation.

The techniques employed to collect and store trematodes vary between research groups, and although these differences are sometimes necessitated by distinctions in the hosts examined, they are more commonly an artefact of instruction. As a general rule, we tend to follow what we were taught rather than explore new techniques. A major reason for this is that there are few technique papers in the published literature. Inspired by a collaborative workshop at the Trematodes 2024 symposium, we outline our techniques and processes for collecting adult trematodes from fishes and discuss the improvements we have made over 40 years of dissections of 20,000+ individual marine fishes. We present these techniques for two reasons: first, to encourage unified methods across the globe, with an aim to produce optimally comparable specimens across temporal periods, across geographic localities, and between research groups; and second, as a resource for inexperienced researchers. We stress the importance of understanding differences in host biology and the expected trematode fauna, which ultimately enables organised and productive dissections. We outline our dissection method for each key organ separately, discuss handling, fixation, and storage methods to generate the most uniform and comparable samples, and explore ethical considerations, issues of accurate host identification, and the importance and potential of clear record keeping.

For many trematode species, individual reproductive parthenitae in first intermediate host colonies senesce, die, and are replaced by newly born parthenitae. The times involved in these processes are poorly understood. Here, we present an approach to estimate parthenita death rates and lifespans that uses readily obtainable data on senescent parthenita frequencies, brood sizes, and offspring (cercaria) release rates. The onset of parthenita senescence is often marked by the degeneration and disappearance of the germinal mass, its source of new offspring. Following germinal mass loss, the remaining viable offspring in a senescent parthenita finish development and are birthed before parthenita death. Therefore, a senescing parthenita’s remaining lifespan is the time it takes for all its viable offspring to mature and exit. We can estimate this time by measuring whole-colony (infected snail) cercaria shed rates, dissecting colonies to count reproductives, and then apply the per redia cercaria production rate to the observed brood sizes of senescent parthenitae. The per-capita parthenita death rate is then calculated as the proportion of parthenitae that are senescent divided by their average remaining lifespan. Reproductive parthenita lifespan is the inverse of this death rate. We demonstrate the approach using philophthalmid trematodes, first providing documentation of a free-floating germinal mass in 4 philophthalmids, and then, for 3 of those species, estimating parthenita senescence rates, death rates, and lifespans. This method should be broadly applicable among trematode species and help inform our understanding of trematode colony dynamics, social structure, and the evolution of parthenita senescence.

The description and delineation of trematode species is a major ongoing task. Across the field there has been, and currently still is, great variation in the standard of this work and in the sophistication of the proposal of taxonomic hypotheses. Although most species are relatively unambiguously distinct from their congeners, many are either morphologically very similar, including the major and rapidly growing component of cryptic species, or are highly variable morphologically despite little to no molecular variation for standard DNA markers. Here we review challenges in species delineation in the context provided to us by the historical literature, and the use of morphological, geographical, host, and molecular data. We observe that there are potential challenges associated with all these information sources. As a result, we encourage careful proposal of taxonomic hypotheses with consideration for underlying species concepts and frank acknowledgement of weaknesses or conflict in the data. It seems clear that there is no single source of data that provides a wholly reliable answer to our taxonomic challenges but that nuanced consideration of information from multiple sources (the ‘integrated approach’) provides the best possibility of developing hypotheses that will stand the test of time.

Over the years, the number of parasitic helminth species discoveries has not ceased to increase and the popularisation of the use of molecular methods has contributed greatly to sustain the growth in knowledge. However, molecular approaches evolved rapidly in the last 20 years. I argue that the research community working on parasitic helminths has lagged behind in the application of molecular methods that examine multiple loci to study species diversity. In this paper, I review the recent historical trends in the molecular markers used to study trematode diversity. Except for the emergence of pioneer mitogenome studies, the use of markers has not changed in the past 10 years. It is still restricted to single locus or a combination of two, rarely three, mitochondrial and ribosomal loci. I identify past and current molecular approaches providing data on multiple loci across the genome which have found resistance in the trematode and the helminth parasitology fields over the last four decades. I discuss how the knowledge gained from the analysis of genome-wide markers would benefit research on parasite diversity today, in particular for cases of species complexes, cryptic (or nearly cryptic) species, recently diverged species, and species with a complex taxonomic history, or a history of suspected mitonuclear discordance as well as for taxa with wide geographical distributions or species with disjoint distributions. Furthermore, I argue that both, studies with classical markers and reduced-representation genome studies providing genome-wide markers should not walk different paths but feedback on each other to advance the field forward. I examine some challenges and make recommendations for obtaining high-throughput molecular data of parasitic helminths.

The freshwater fish fauna of southern Africa is highly diverse; however, the magnitude of parasitic species they host is unevenly known. The region’s documented adult trematode fish fauna is sparse, while the opposite is evident for intermediate trematode stages. Perceived difficulty in identification of underdeveloped stages lead to the exclusion of reporting metacercariae or lack either morphological or molecular data resulting in a depauperate comparative molecular data repository for species of the region and Africa as a whole. In an effort to address the morphological and molecular data void of the parasite fauna of southern African freshwater fishes, we sought to comprehensively investigate and characterise this fauna. Here we report on three metacercarial forms of Clinostomum (Clinostomidae) from three fish families (Clariidae, Mochokidae, and Mormyridae), provide the first report of a species of the Cryptogonimidae from a cyprinid host in South Africa, and include molecular data for the partial 28S rDNA, ITS1–2 and COI mtDNA regions of these metacercarial forms. Our clinostomid specimens morphologically and genetically corresponded with Clinostomum brieni (e.g., Clarias gariepinus) and Clinostomum ‘morphotype 2’ and ‘morphotype 3’ per Caffara et al. (2017) from the mormyrid Marcusenius pongolensis and the mochokid catfish Chiloglanis sp., respectively. Our cryptogonimid metacercariae did not correspond with any known species or available molecular sequence data; however, the presence of robust circumoral spines on the oral sucker indicated that they are either a species of Acanthostomum or Proctocaecum. The molecular data we provide are the first for an Acanthostomum/Proctocaecum-type cryptogonimid from Africa.