A survey of nocturnality and risk for savanna chimpanzees at Assirik, Senegal.

Chimpanzees (Pan troglodytes) in Senegal may use nocturnality to mitigate hyperthermia risk in semi-arid environments but the degree of nocturnality for such chimpanzees also in sympatry with large carnivores remains uncertain. We compared diel activity among chimpanzees and their potential predators at Assirik in Niokolo-Koba National Park and contextualized these findings relative to other unit-groups in savanna landscapes. From 2015-2018, we generated a predator inventory using multi-modal methods and monitored the diel activity of chimpanzees and predators with camera traps [ N = 2092 camera trap (CT) days]. From 2015-2023, we also surveyed for evidence of predation during recce walks. Six potential nonhuman predators occur at Assirik, including lions (Panthera leo), leopards (Panthera pardus), spotted hyenas (Crocuta crocuta), African wild dogs (Lycaon pictus), Nile crocodiles (Crocodylus niloticus), and rock pythons (Python sebae). We documented one suspected case of a predator killing a chimpanzee. Nocturnality comprised 12.7% of CT events for chimpanzees and these events were more concentrated at twilight. Chimpanzees were more active during the day, predators were more active at night, and there was substantial temporal overlap among chimpanzees and potential predators during twilight intervals. Our findings support the hypothesis that savanna chimpanzees in Senegal are active at night in response to the extremely hot environment. We hypothesize that Assirik chimpanzees experience a tension between decreasing hyperthermia and increasing predation risk during nocturnality.

Updating Chimpanzee Nesting Data at Mount Assirik (Niokolo Koba National Park, Senegal): Implications for Conservation.

The Niokolo Koba National Park (NKNP) is the largest protected area in Senegal and lies at the northern limit of the chimpanzee's range in West Africa. Recent information on nesting behavior and factors influencing nesting behavior is available for several sites outside NKNP. However, the information available for NKNP is obsolete. Considering that the adequate management of chimpanzee populations cannot be achieved without strong scientific knowledge, it is essential to update data on chimpanzee, Pan troglodytes verus, nesting behavior in NKNP. For this reason, we surveyed their habitat in Mt. Assirik and recorded 626 chimpanzee nests. The results of the study showed that chimpanzees nest more often in closed-canopy habitats such as gallery forests. The average nest height observed in this study was 8.07 ± 0.36 m, varying between 2 and 20 m, which is well below the heights described in most sites where chimpanzees cohabit with large carnivores. Botanical surveys confirmed that chimpanzees select tree species bearing their nests. In Assirik, 12 of the 37 tree species bearing nests are the most used. The nest decay rate (or the time it takes for a nest to go from the fresh to the skeletal stage) at Assirik averaged 68.8 ± 5.8 days.

Reciprocity and beyond: Explaining meat transfers in savanna-dwelling chimpanzees at Fongoli, Senegal.

OBJECTIVES: To understand the function of food sharing among our early hominin ancestors, we can turn to our nonhuman primate relatives for insight. Here, we examined the function of meat sharing by Fongoli chimpanzees, a community of western chimpanzees (Pan troglodytes verus) in southeastern Sénégal. MATERIALS AND METHODS: We tested three non-mutually exclusive hypotheses that have been used to explain patterns of food sharing: kin selection, generalized reciprocity, and meat-for-mating opportunities. We analyzed meat sharing events (n = 484) resulting from hunts, along with data on copulations, age-sex class, and kinship to determine which variables predict the likelihood of meat sharing during this study period (2006-2019). RESULTS: We found full or partial support for kin selection, direct reciprocity, and meat-for-mating-opportunities. However, the analyses reveal that reciprocity and a mother/offspring relationship were the strongest predictors of whether or not an individual shared meat. CONCLUSIONS: The results of this study emphasize the complexity of chimpanzee meat sharing behaviors, especially at a site where social tolerance offers increased opportunities for meat sharing by individuals other than dominant males. These findings can be placed in a referential model to inform hypotheses about the sensitivity of food sharing to environmental pressures, such as resource scarcity in savanna landscapes.

Gastrointestinal parasites of baboons (Papio papio) in Niokolo-Koba National Park, Senegal.

Background: Primates can harbor parasites that could be pathogenic or not for humans and primates themselves. It is necessary to know the parasitological situation of the primates that are under surveillance in the park. Aim: To estimate the prevalence and diversity of gastrointestinal parasites, including zoonotic potential parasites, in baboons in the Niokolo-Koba National Park located in Senegal. Method: Fecal samples (n = 50) from two groups of baboons (A and B) were collected in October 2019. The samples were processed using the flotation technique and the modified Ritchie method. Slides were examined microscopically and the parasite identification was based on morphology, color, and parasite content. Results: A total of seven nematodes (Strongyloides sp., Trichirus sp., Ancylostoma sp., Mammo monogamus, Enterobius sp., Strongyloides stercoralis, Strongyle digestif), one cestode (Bertiella sp.), and one trematode (Fasciolopsis sp.) were identified. The overall prevalence was 78%, while the prevalence of poly-infected samples was 49%. The parasite with zoonotic potential, S. stercoralis, was identified in group B samples. Trichuris sp., which is common and pathogenic to humans and primates, was present with prevalence of 52% and of 32% in groups A and B, respectively. Conclusion: These results suggest that baboons are infested with zoonotic parasites and this situation could expose people working in this park to infection. Contact between humans and these baboons or their feces could expose them to infection with zoonotic parasites.

Chimpanzees (Pan troglodytes) in savanna landscapes.

Chimpanzees (Pan troglodytes) are the only great apes that inhabit hot, dry, and open savannas. We review the environmental pressures of savannas on chimpanzees, such as food and water scarcity, and the evidence for chimpanzees' behavioral responses to these landscapes. In our analysis, savannas were generally associated with low chimpanzee population densities and large home ranges. In addition, thermoregulatory behaviors that likely reduce hyperthermia risk, such as cave use, were frequently observed in the hottest and driest savanna landscapes. We hypothesize that such responses are evidence of a "savanna landscape effect" in chimpanzees and offer pathways for future research to understand its evolutionary processes and mechanisms. We conclude by discussing the significance of research on savanna chimpanzees to modeling the evolution of early hominin traits and informing conservation programs for these endangered apes.

A comparative study of the ultrastructural characteristics of the mature spermatozoa of two fellodistomids Tergestia clonacantha and T. laticollis and contribution to the phylogenetic knowledge of the Gymnophalloidea.

The ultrastructure of the mature spermatozoa of Tergestia clonacantha and T. laticollis collected from the digestive tracts of fishes from New Caledonia is described using transmission electron microscopy and compared to that of related species. The spermatozoa of the two species exhibit the general pattern described in most digeneans, namely two axonemes with the 9 + "1" pattern of the Trepaxonemata, nucleus, mitochondrion, cortical microtubules, an external ornamentation of the plasma membrane, spine-like bodies and granules of glycogen. The spermatozoa of T. clonacantha and T. laticollis show the same ultrastructural model with some specificities in each case, particularly in the disposition of the structures in the posterior extremities of the spermatozoon. This study confirms that ultrastructural characters of the mature spermatozoon are useful tools for the phylogenetic analysis of the Digenea.

TITLE: Étude comparative des caractéristiques ultrastructurales des spermatozoïdes mûrs de deux Fellodistomidae, Tergestia clonacantha et T. laticollis, et contribution à la connaissance phylogénétique des Gymnophalloidea. ABSTRACT: L'ultrastructure des spermatozoïdes mûrs de Tergestia clonacantha et T. laticollis, prélevés dans le tube digestif de poissons de Nouvelle-Calédonie, est décrite par microscopie électronique à transmission et comparée à celle d'espèces apparentées. Les spermatozoïdes des deux espèces présentent la structure générale décrite chez la plupart des digènes, à savoir deux axonèmes du type 9 + « 1 ¼ des Trepaxonemata, un noyau, une mitochondrie, des microtubules corticaux, des ornementations externes de la membrane plasmique, des corps épineux et des granules de glycogène. Les spermatozoïdes de T. clonacantha et T. laticollis présentent le même modèle ultrastructural avec quelques spécificités dans chaque cas, notamment dans la disposition des structures aux extrémités postérieures du spermatozoïde. Cette étude confirme que les caractères ultrastructuraux du spermatozoïde mûrs sont des outils utiles pour l'analyse phylogénétique des Digenea.

Spermatological characteristics of Sclerodistomoides pacificus (Digenea, Sclerodistomoididae) a parasite of the flying fish Cheilopogon pinnatibarbatus (Teleostei, Exocoetidae).

Sclerodistomoides pacificus is the only species described now in Sclerodistomoididae. We present in this paper the first ultrastructural data of the mature spermatozoon of a species from the genus Sclerodistomoides. Adult specimens of S. pacificus (Digenea: Hemiuroidea: Sclerodistomoididae), were parasites of the gall-bladder of the teleost fish Cheilopogon pinnatibarbatus captured in the Atlantic Ocean, near Dakar (Senegal). The male gamete is a filiform cell which exhibits a similar ultrastructural organization to that reported in most species belonging to the Hemiuroidea with two axonemes of the 9 + '1' pattern of trepaxonematans, a nucleus, a mitochondrion, external ornamentation of the plasma membrane not associated with cortical microtubules and located in the anterior region of the spermatozoon, and parallel cortical microtubules disposed in one side of the spermatozoon. However, the present study allowed describing for the first time a moniliform mitochondrion in the Hemiuroidea. The presence of a moniliform mitochondrion and the absence of filamentous external ornamentation described in other Hemiuridae: Lecithochirium microstomum, L. musculus and Hemiurus appendiculatus are a good tool for phylogenetic purposes in the Hemiuroidea. Moreover, spermatological organisation and model are discussed in context with those of previous studies in the Hemiuroidea.

Towards systematic and evidence-based conservation planning for western chimpanzees.

As animal populations continue to decline, frequently driven by large-scale land-use change, there is a critical need for improved environmental planning. While data-driven spatial planning is widely applied in conservation, as of yet it is rarely used for primates. The western chimpanzee (Pan troglodytes verus) declined by 80% within 24 years and was uplisted to Critically Endangered by the IUCN Red List of Threatened Species in 2016. To support conservation planning for western chimpanzees, we systematically identified geographic areas important for this taxon. We based our analysis on a previously published data set of modeled density distribution and on several scenarios that accounted for different spatial scales and conservation targets. Across all scenarios, typically less than one-third of areas we identified as important are currently designated as high-level protected areas (i.e., national park or IUCN category I or II). For example, in the scenario for protecting 50% of all chimpanzees remaining in West Africa (i.e., approximately 26,500 chimpanzees), an area of approximately 60,000 km2 was selected (i.e., approximately 12% of the geographic range), only 24% of which is currently designated as protected areas. The derived maps can be used to inform the geographic prioritization of conservation interventions, including protected area expansion, "no-go-zones" for industry and infrastructure, and conservation sites outside the protected area network. Environmental guidelines by major institutions funding infrastructure and resource extraction projects explicitly require corporations to minimize the negative impact on great apes. Therefore, our results can inform avoidance and mitigation measures during the planning phases of such projects. This study was designed to inform future stakeholder consultation processes that could ultimately integrate the conservation of western chimpanzees with national land-use priorities. Our approach may help in promoting similar work for other primate taxa to inform systematic conservation planning in times of growing threats.

Informing Protection Efforts for Critically Endangered Chimpanzees (Pan troglodytes verus) and Sympatric Mammals amidst Rapid Growth of Extractive Industries in Senegal.

Updated information on Critically Endangered western chimpanzees (Pan troglodytes verus) in Senegal is urgently needed, given that gold mining is rapidly transforming landscapes and livelihoods. Specifically, biodiversity assessments will better elucidate the chimpanzee extinction risk here and yield baselines for monitoring. We compared mammal species richness between Fongoli (unprotected) and Assirik in Niokolo-Koba National Park to assess the efficacy of the only nationally protected area where chimpanzees range in this country. The primary habitat difference between these sites was the degree of human activity. Although Assirik and Fongoli had similar mammal assemblages and were equivalent in primate species richness, the protected area was higher in species richness overall, particularly for ungulates and carnivores. The protected status and management plan of Niokolo-Koba almost certainly resulted in fewer mammal extinctions. In unprotected areas, the flexible behavioural responses of chimpanzees to human-driven landscape dynamics is likely essential to their long-term survival. Furthermore, the near absence of chimpanzee killings in this country greatly facilitates conservation efforts. Given that mineral extraction is a key development strategy in Senegal, we encourage primatologists to collaborate with mining stakeholders to ensure that sustainability programming includes contributions from experts in savannah chimpanzee behavior and ecology.

Ultrastructure of mature spermatozoa of three Bucephalidae (Prosorhynchus longisaccatus, Rhipidocotyle khalili and Bucephalus margaritae) and phylogenetic implications.

We describe here the mature spermatozoa of three species of bucephalids, namely Bucephalus margaritae, Rhipidocotyle khalili and Prosorhynchus longisaccatus. This study provides the first ultrastructural data on the genera Bucephalus and Rhipidocotyle and enabled us to confirm the model of the mature spermatozoon in the Bucephalinae. The spermatozoon exhibits two axonemes with the 9 + "1" pattern of the Trepaxonemata, one of which is very short, lateral expansion, external ornamentation of the plasma membrane located in the anterior extremity of the spermatozoon and associated with cortical microtubules, spine-like bodies, a mitochondrion, and a nucleus. The maximum number of cortical microtubules is located in the anterior part of the spermatozoon. However, more studies are needed to elucidate if spine-like bodies are present in all the Bucephalinae or not. In the Prosorhynchinae, the mature spermatozoon exhibits a similar ultrastructural pattern. Some differences are observed, particularly the axoneme lengths and the arrangement of the spine-like bodies. The posterior extremity of the spermatozoon in the Bucephalinae exhibits only the nucleus, but prosorhynchines have microtubules.

Nest-Building Behaviour by Chimpanzees (Pan troglodytes verus) in the Non-Protected Area of Diaguiri (Kedougou, Senegal): Implications for Conservation.

Diaguiri is a non-protected area situated in south-eastern Senegal. Chimpanzees (Pan troglodytes verus) have been living here since 1999 but information on the characteristics of their nest sites have been lacking until now. Furthermore, we implemented nest survey methods recently standardised to arid landscapes during a national chimpanzee survey. Variations in nest decay rates are poorly understood in savanna-woodland mosaic habitats. This study aims to describe in Diaguiri: (1) the distribution of chimpanzee nests and the location of their core sleeping area, (2) the tree species used for nest building, (3) the association between nests and nesting tree heights, and (4) differences in nest decay rates between habitats. We surveyed 43 km of chimpanzee nesting habitats and recorded 871 nests in gallery forest and woodland habitats between March 2015 and June 2016. Diaguiri chimpanzees used at least 17 tree species for nest building, and 58.8% of nests were in only 3 tree species: Pterocarpus erinaceus, Anogeissus leiocarpus and Diospyros mespiliformis. Nest decay rate was faster in gallery forests than in woodlands. These data are of great value for understanding habitat use by a Critically Endangered subspecies of savanna chimpanzee, for population density estimates of the species and for conservation programmes in this region and in savanna biomes.

Ultrastructural characteristics of the mature spermatozoon of the digenean Sclerodistomum italicum (Stossich, 1893) (Hemiuroidea, Sclerodistomidae) intestinal parasite of Hypocanthus amia (Teleostei, Carangidae).

The mature spermatozoon of Sclerodistomum italicum is filiform, tapered at both ends and shows the following features: 2 axonemes of the 9+"1" pattern of the Trepaxonemata, mitochondrion, nucleus and parallel cortical microtubules. The specific features of the spermatozoon of S. italicum include the simultaneous presence of two types of extramembranous ornamentations, the presence of short cortical microtubules in the anterior part of the spermatozoon and the presence of only one bundle of cortical microtubules in the median part of the spermatozoon. Thus far these structures are known only in the Hemiuroidea. The presence of filamentous ornamentation in the anterior extremity of the spermatozoon has not previously been described in the Sclerodistomidae. Similar to spermatozoa of other hemiuroideans, S. italicum lack spine-like bodies described in spermatozoa of many digenean taxa. The posterior extremity of the spermatozoon exhibits the same ultrastructural characteristics typical of the Hemiuroidea.

Ultrastructural study of the spermatozoon of Hemiurus appendiculatus (Digenea, Hemiuroidea, Hemiuridae), a parasite of Boops boops (Pisces, Teleostei, Sparidae) off Senegal.

The mature spermatozoon of Hemiurus appendiculatus exhibits the general pattern described in most of the digenean namely, two axonemes of the 9 + "1" pattern of the Trepaxonemata, a nucleus, a filiform mitochondrion, external ornamentations of the plasma membrane and parallel cortical microtubules located on one side of the spermatozoon. In this study, we show for the second time in a digenean spermatozoon the presence of microtubules of the second axoneme associated each in a short length with an external ornamentation, seven cortical microtubules, a terminal bulge in the anterior spermatozoon extremity separate from the remainder of the spermatozoon by a transverse constriction and the simultaneous presence of the external ornamentation of the plasma membrane with a filamentous ornamentation. The external ornamentations have a differentiated distribution, cover almost the anterior region of the spermatozoon, are more abundant around the first axoneme and extend backwards over a short distance around the second axoneme. This study also allowed us to reveal for the first time the existence in the Hemiuroidea of an axoneme that begins to disrupt before reaching the front end of the mitochondrion.

Ultrastructural characters of the spermatozoa in Digeneans of the genus Bianium Stunkard, 1930 (Digenea, Lepocreadiidae) parasites of fishes: a comparative study of Bianium plicitum and Bianium arabicum.

We present in this paper the first ultrastructural study of the spermatozoon of species of the genus Bianium Stunkard, 1930, namely Bianium plicitum and Bianium arabicum belonging to the family Lepocreadiidae. Specimens are parasites of the tetraodontid teleosts Lagocephalus laevigatus caught in the Atlantic Ocean and Lagocephalus sceleratus caught in Pacific Ocean, respectively. Live worms were fixed and processed for transmission electron microscopy. Observations of numerous cross and longitudinal sections of the spermatozoa of B. plicitum and B. arabicum demonstrated the general pattern described in all previously studied Lepocreadioidea, namely two axonemes with 9+ "1" pattern of Trepaxonemata, two mitochondria, a nucleus, parallel cortical microtubules, and an external ornamentation of the plasma membrane associated with spine-like bodies. The maximum number of cortical microtubules and the external ornamentation were not located at the anterior extremity of the spermatozoon but at a more posterior level, as in all the species of Lepocreadioidea studied to date. Differences observed between spermatozoa of B. plicitum and B. arabicum were observed in: (1) the location of the first mitochondrion, (2) size of cortical microtubules which were longer in B. arabicum than in B. plicitum, and (3) the order of the disappearance of the axonemes, mitochondrion, and cortical microtubules toward the posterior region of the spermatozoon. Only the nucleus reaches into the posterior extremity of the spermatozoon in these two species. The general pattern of the spermatozoon described in these two species exhibit the general model in the genus Bianium.

Ultrastructural characters of the spermatozoa in Digeneans of the genus Lecithochirium Lühe, 1901 (Digenea, Hemiuridae), parasites of fishes: comparative study of L. microstomum and L. musculus.

This study provides the first ultrastructural data of spermatozoa in the genus Lecithochirium. The spermatozoa of L. microstomum (from Trichiurus lepturus in Senegal) and L. musculus (from Anguilla anguilla in Corsica) exhibit the general pattern described in the great majority of the Digenea, namely two axonemes with the 9 + "1" pattern typical of the Trepaxonemata, one mitochondrion, a nucleus, parallel cortical microtubules and external ornamentation of the plasma membrane. Spermatozoa of L. microstomum and L. musculus have some specific features such as the presence of a reduced number of cortical microtubules arranged on only one side of the spermatozoon, the lack of spine-like bodies and expansion of the plasma membrane. The external ornamentation of the plasma membrane entirely covers the anterior extremity of the spermatozoa. The ultrastructure of the posterior extremity of the spermatozoa corresponds to the pattern previously described in the Hemiuridae, characterized by only singlets of the second axoneme. A particularity of these spermatozoa is the organization of the microtubule doublets of the second axoneme around the nucleus in the posterior part of the spermatozoon.

Vitellogenesis of the digenean Plagiorchis elegans (Rudolphi, 1802) (Plagiorchioidea, Plagiorchiidae).

The ultrastructural organization of vitellogenesis of Plagiorchis elegans (Rudolphi, 1802), experimentally obtained from the golden hamster Mesocricetus auratus (Linnaeus, 1758), is described using transmission electron microscopy. This study is the first ultrastructural study of vitellogenesis in a member of the superfamily Plagiorchioidea. The four stages usually observed during vitellogenesis are described: stage I, cytoplasm of the vitellocytes mainly filled with ribosomes and few mitochondria; stage II, beginning of the synthetic activity; stage III, active synthesis of the shell globule clusters; stage IV, vitellocytes are filled with shell globule clusters and contain several lipid droplets, and glycogen granules are grouped around clusters and droplets. Vitellogenesis in P. elegans is compared with that of other Digenea. The differences among P. elegans and previously studied digeneans include, but are not limited to the occurrence of dense coiled endoplasmic reticulum saccules and the concentration of glycogen in the mesenchyme, which may be considered as a fifth stage of maturation of the vitelline glands. This peculiarity was not observed in all trematodes, which clearly indicates differences in the vitellogenesis in various digenean lineages at different stages of maturation of their vitelline cells.

Morphometrical and genetic comparison of two nematode species: H. spumosa and H. dahomensis (Nematoda, Heterakidae).

Heterakis is a genus of parasitic nematodes, the majority of which are found in ground-feeding birds and only rarely in mammals. The best-known species is Heterakis spumosa, a parasite associated with the cosmopolitan invasive rodent Rattus rattus of Asiatic origin. Heterakis dahomensis was described in 1911 as a parasite of the Gambian giant rat (Cricetomys gambianus) from Benin (Africa), subsequently synonymized to H. spumosa by Hall (1916). The study of helminths in African rodents is scarce and patchy. Since the original description of H. dahomensis, there have been only a few reports from Africa of species belonging to the genus Heterakis and the validity of this species has never in fact been confirmed or rejected. In the present study individual Heterakis spp. were collected from C. gambianus from Senegal. The morphological data taken point to differences between Heterakis dahomensis and H. spumosa, specifically in the number of tail papillae in males and in the vulva cuticular processes of females. In addition, molecular data revealed differences between these taxa and so H. dahomensis should be considered as a valid species. Moreover, recent changes in the systematics of the genus Cricetomys mean that it is now necessary to study the morphology and genetics of the Heterakis specimens collected from Cricetomys spp. (previously assigned to C. gambianus) in order to determine their taxonomic status as either H. dahomensis o H. spumosa.

Ultrastructure of the spermatozoon of the digenean Plagiorchis elegans (Rudolphi, 1802) (Plagiorchioidea, Plagiorchiidae).

The ultrastructure of the mature spermatozoon of the type genus of the Plagiorchiidae Plagiorchis elegans (Rudolphi, 1802), a parasite of the Golden hamster, Mesocricetus auratus is described. This study is the first ultrastructural study of the spermatozoon of a Plagiorchis, the second of a plagiorchiid species and only the third in the Plagiorchioidea. Previously data on spermatozoon ultrastructure existed only for the plagiorchiid Enodiotrema reductum and the omphalometrid Rubenstrema exasperatum. The mature spermatozoon of P. elegans exhibited the general pattern described in most digenean species, namely two axonemes of the 9 + "1" Trepaxonemata pattern, nucleus, mitochondria, external ornamentation of the plasma membrane, spine-like bodies, and glycogen granules. However, the rather typical expansion of the plasma membrane is not found in P. elegans. Another peculiarity of the spermatozoon of P. elegans is the presence of a structure called thin cytoplasm termination. Spermatozoon ultrastructure of P. elegans is compared with that of E. reductum and R. exasperatum. Spermatozoon of P. elegans conforms to the general pattern described in E. reductum. Thus, this study further expands our knowledge on the spermatozoon ultrastructure among the members of the Plagiorchioidea, one of the most phylogenetically derived groups of the digenea.

Ultrastructure of the spermatozoon of the digenean Lecithocladium excisum (Rudolphi, 1819) (Hemiuroidea: Hemiuridae), a parasite of marine teleosts in Senegal.

The present study describes the ultrastructure of the mature spermatozoon of Lecithocladium excisum (Rudolphi, 1819) (Digenea: Hemiuroidea: Hemiuridae) from the stomach of the marine teleost Scomber japonicus Houttuyn (Scombridae) captured in the Atlantic Ocean, off Dakar (Senegal). The ultrastructural organization of the spermatozoon of L. excisum follows the general model described in most digeneans. It presents two axonemes of the 9+'1' pattern of the Trepaxonemata, nucleus, mitochondrion and parallel cortical microtubules, among other characters. However, some particularities of the spermatozoon of L. excisum are (i) the presence of a membranous ornamentation not associated with cortical microtubules in its anterior extremity, (ii) the presence of a very reduced number of cortical microtubules located only in the ventral side of the spermatozoon and (iii) the absence of several structures described in most digeneans such as spine-like bodies and cytoplasmic expansions.

Spermiogenesis and the spermatozoon ultrastructure of Robphildollfusium fractum (Digenea: Gyliauchenidae), an intestinal parasite of Sarpa salpa (Pisces: Teleostei).

Spermiogenesis in Robphildollfusium fractum begins with the formation of a differentiation zone containing: two centrioles, each bearing striated rootlets, nucleus, several mitochondria and an intercentriolar body constituted by seven electron-dense layers. The two centrioles originate two free flagella growing orthogonally to the median cytoplasmic process. Later, the free flagella rotate and undergo proximodistal fusion with the median cytoplasmic process. Nuclear and mitochondrial migrations occur before this proximodistal fusion. Finally, the young spermatozoon detaches from the residual cytoplasm after the constriction of the ring of arched membranes. The spermatozoon of R. fractum exhibits two axonemes of different length of the 9+'1' trepaxonematan pattern, nucleus, two mitochondria, two bundles of parallel cortical microtubules, external ornamentation of the plasma membrane, spine-like bodies and granules of glycogen. Additionally, a shorter axoneme, which does not reach the nuclear region, the presence of an electron-dense material in the anterior spermatozoon extremity and the morphologies of both spermatozoon extremities characterize the mature sperm of R. fractum.