NEW SPECIES OF ALIPPISTRONGYLUS (TRICHOSTRONGYLINA: NIPPOSTRONGYLINAE) FROM THE ELEGANT RICE RAT, EURYORYZOMYS NITIDUS, OF LA CONVENCIÓN, PERU.

A new species of AlippistrongylusDigiani and Kinsella, 2014, was found in the intestines of the elegant rice rat, Euryoryzomys nitidus, collected in the Amazon rainforest. These mammals were preserved in alcohol and archived in the Colección Científica de Mamíferos del Museo de Historia Natural de la Universidad Mayor de San Marcos and accessed for the observation and investigation for the diversity of their endo-parasites. A total of 857 worms were recovered from the 3 individuals examined. Morphological investigations and comparisons with the only known species of the genus indicate that this is an undescribed species. These nematodes feature the diagnostic characteristic of the genus, being a bifurcated posterior end that consists of a tail and a conical appendage near the level of the vulva and uninterrupted ridges in the synlophe of unequal size. However, the orientation of this conical appendage on the female tail, features of the synlophe, and shape of the copulatory bursa warrant the proposal of an amended diagnosis to include character variability detected in the new species.

Morphological and Molecular Data Reveal Two New Species of Viannaia (Nematoda: Viannaiidae), Parasitizing Opossums (Mammalia: Didelphidae) in Mexico.

Two new species of Viannaia from the intestine of the North American opossums, Didelphis virginiana (Virginia opossum), and Philander opossum (gray four-eyed opossum), are described based on morphological and molecular data, through an integrative taxonomic approach. Maximum likelihood and Bayesian inference analyses for each dataset and the concatenated dataset were performed using a mitochondrial cytochrome c oxidase subunit 1 (COI) gene, and the nuclear ribosomal internal transcribed spacer region (ITS1-5.8S-ITS2). The phylogenetic analyses revealed 2 new species that occur in Mexico, one from the western state of Colima and another from the southern state of Chiapas. Our phylogenetic trees for both molecular markers and concatenated datasets yielded similar topologies with high bootstrap values and posterior probabilities. Viannaia is recovered as a monophyletic group, but the family Viannaiidae appears as non-monophyletic, due to the position of Travassostrongylus scheibelorum, similar to previous studies. Finally, the morphology of Viannaia and Hoineffia is discussed.

A New Species of Bidigiticauda (Nematoda: Strongylida) from the Bat Artibeus Planirostris (Chiroptera: Phyllostomidae) in the Atlantic Forest and a Molecular Phylogeny of the Molineid Bat Parasites.

The nematode genus Bidigiticauda has 2 species (Bidigiticauda vivipara and Bidigiticauda embryophilum), which are parasites of bats from the Neotropical region. The present paper describes a new species of Bidigiticauda from a male Artibeus planirostris specimen collected in the Pratigi Environmental Protection Area in Bahia state, Brazil. The new species, Bidigiticauda serrafreirei n. sp., differs from B. embryophilum by having longer spicules, rays 5 and 6 arising from a common trunk and bifurcating in its first third, rays 3 and 4 emerging slightly separated from each other, and dorsal rays reaching the margin of the caudal bursa. The new species also differs from B. vivipara by the dorsal ray bifurcating at the extremity of the trunk. A molecular phylogenetic analysis was conducted to determine the evolutionary affinities of Bidigiticauda serrafreirei n. sp. within the Strongylida, which identified a clade that grouped Bidigiticauda with the other members of the Anoplostrongylinae. However, the molineid subfamilies did not group together, indicating that the family Molineidae is polyphyletic. Further analyses, which include additional taxa and genetic markers, should elucidate the complex relationships within the Molineidae, in particular its subfamilies and the evolution of the traits that define these groups.

Zygocaulus nagoensis n. gen. and n. sp. (Trichostrongyloidea: Dictyocaulidae), a Peculiar Bursate Nematode Collected from an Alien Frog, Polypedates leucomystax, in Nago, Okinawa Island, Japan.

A peculiar bursate nematode, Zygocaulus nagoensis n. gen. and n. sp. (Trichostrongyloidea: Dictyocaulidae: Mertensinematinae), was described from an alien frog, Polypedates leucomystax (Anura: Rhacophoridae), collected on Okinawa Island, Japan. It is related to Mertensinema and Borrellostrongylus, the only hitherto known genera of Mertensinematinae, but is readily distinguished from them by having only 2 pairs of lateral rays and simple distal ends of the dorsal ray branches.

A new heligmonellid (Nematoda) species of the genus Stilestrongylus in Euryoryzomys russatus (Rodentia: Sigmodontinae) in the Atlantic Forest, southern Brazil.

A new species of Nippostrongylinae (Nematoda: Heligmonellidae), Stilestrongylus rolandoi n. sp., is described from specimens collected from the small intestine of the rodent Euryoryzomys russatus in the Atlantic Forest (Santo Amaro da Imperatriz, Santa Catarina state, southern Brazil). The genus Stilestrongylus includes 23 species, which parasitize rodents occurring in the Neotropical region. Stilestrongylus aureus (Durette-Desset & Sutton, 1985) from Argentina, S. azarai (Durette-Desset & Sutton, 1985) from Argentina, S. flavescens (Sutton & Durette-Desset, 1991) from Uruguay, S. franciscanus (Digiani & Durette-Desset, 2002) from Argentina, S. gracielae (Digiani & Durette-Desset, 2006) from Argentina, and S. oryzomysi (Sutton & Durette-Desset, 1991) from Argentina are closely related to Stilestrongylus rolandoi n. sp., all having caudal bursa patterns of types 1-4 in one of the lobes. Stilestrongylus rolandoi n. sp. is distinguished from the aforementioned species by its ray 6 being short in relation to rays 4 and 5, which are long and robust, and by having caudal bursa patterns of types 1-4 in both lobes. The new species has 27 ridges in the mid-body in males, and 24 in females, and has one of the highest ratios of spicule length to body length (21-33%) in this genus.

Molecular marker sequences of cattle Cooperia species identify Cooperia spatulata as a morphotype of Cooperia punctata.

The genus Cooperia includes important parasites of ruminants and currently contains 34 accepted species. However, even for those species infecting livestock, there is a considerable lack of molecular information and many species are only identifiable using subtle morphological traits. The present study aimed to provide molecular data to allow diagnosis of Cooperia species infecting cattle. Partial sequences of two mitochondrial (cytochrome oxidase 2, 12S rRNA gene) and two nuclear genes (isotype 1 ß tubulin gene including two introns, internal transcribed spacers (ITS) were obtained from morphologically identified specimens of Cooperia pectinata, Cooperia punctata and Cooperia spatulata as well as from larvae of pure Cooperia oncophora and C. punctata laboratory isolates. Pairwise identity of ITS-2 sequences was very high and it was the only region able to identify a specimen as Cooperia sp. However, the ITS-2 was unreliable for diagnosis at the species level. All other marker sequences could not unequivocally be allocated to the genus Cooperia but allowed clear species identification with the exception of the pair C. punctata/C. spatulata for which no significant differences were found for any marker sequence. Maximum-likelihood phylogenetic analyses of individual genes as well as a multi-locus analysis covering all four sequences confirmed that specimen identified as C. spatulata were randomly distributed throughout the C. punctata cluster and formed no group of their own. In contrast, the other Cooperia species formed clearly separated and statistically supported clusters. These data indicate that C. spatulata is most likely only a morphotype of C. punctata and the name should be considered a synonym. Combinations of nuclear and mitochondrial markers should be used to identify morphotypes or cryptic species to benefit from excellent barcoding properties of the latter but allowing proper phylogenetic analyses and controlling for lineage sorting that might occur for mitochondrial genotypes within a species.

A new species of Stilestrongylus (Nematoda, Heligmonellidae) from the Atlantic Forest of Misiones, Argentina, parasitic in Euryoryzomys russatus (Cricetidae, Sigmodontinae).

A new species of Heligmonellidae (Trichostrongylina, Heligmosomoidea), Stilestrongylus kaaguyporai n. sp. is described from the small intestine of Euryoryzomys russatus (Rodentia, Cricetidae, Sigmodontinae) from the Argentine Atlantic Forest, in the Misiones province. The new species was found at Campo Anexo Manuel Belgrano, Reserva de Vida Silvestre Urugua-í and Parque Provincial Urugua-í, with a prevalence of 73% in 15 hosts examined. Stilestrongylus includes 24 Neotropical species, all parasitic in rodents, mostly Sigmodontinae. Stilestrongylus kaaguyporai n. sp. can be differentiated from its congeners by the following characters: caudal bursa dissymmetrical with right lobe larger and pattern of type1-4 in both lobes, rays 6 not forming a lateral trident with rays 4 and 5, rays 8 with dissymmetrical pathway, genital cone hypertrophied with a conspicuous hood-like projection and females with a marked dorso-ventral torsion of the posterior end. This report is the second record of a Stilestrongylus species in E. russatus, increasing to nine the number of parasitic species known from this host.

Pudicinae (Nematoda: Heligmonellidae) Parasitic in Endemic Chilean Rodents (Caviomorpha: Octodontidae and Abrocomidae): Description of a New Species and Emended Description of Pudica degusi (Babero and Cattan) n. comb.

We report the finding of 2 species of Pudica (Nematoda: Heligmonellidae: Pudicinae) in 2 rodents endemic to Chile, the common degu Octodon degus (Octodontidae) and the Bennett's chinchilla rat Abrocoma bennettii (Abrocomidae). Pudica degusi ( Babero and Cattan, 1975 ) n. comb., originally described as a species of Longistriata (Heligmosomidae), was found in the common degu; through the study of its synlophe, the species is reassigned to the Heligmonellidae: Pudicinae and the genus Pudica, and it is revalidated through comparison with the remaining species of the genus. Pudica cattani n. sp. is described from both O. degus and A. bennettii. It is characterized by its large body size, bursal pattern of type 1-3-1 on right lobe, 1-3-1 tending to 1-4 on left lobe, synlophe with 11 ridges including a careen, dorsal ray of the bursa dividing proximally and bursal rays 9 and 10 relatively short. Pudica degusi n. comb. and Pudica cattani n. sp. were found in the same host species but not as coparasitic in the same individuals. The common degu is confirmed as the sole and primary host of Pudica degusi n. comb. It is unlikely that it is the primary host for Pudica cattani n. sp., whose host affinities are less clear mainly due to the scarcity of data. Pudica cattani n. sp. is the first helminth reported from the Bennett's chinchilla rat. Both findings enlarge the host range of the Pudicinae to the families Octodontidae and Abrocomidae, i.e., 9 out of the 11 extant families of caviomorphs, thereby establishing the presence of this nematode subfamily as typical parasites of the Neotropical Hystricognathi.

Ashworthius sidemi Schulz, 1933 and Haemonchus contortus (Rudolphi, 1803) in cervids in France: integrative approach for species identification.

Among gastro-intestinal nematodes, the blood-sucking worms belonging to the subfamily of Haemonchinae are considered to be of pathogenic and economic great importance, particularly in small ruminants. Haemonchus contortus, primary found in domestic ruminants and wild bovines (Mouflon, Chamois), is probably the most studied, but occurrence of Ashworthius sidemi has gradually increased over recent years, especially in Cervids and free roaming wild bovid as the European bison in eastern Europe, and some cases of co-infestation were recently observed on five Roe deer (Capreolus capreolus) and one Red deer (Cervus elaphus) in France. If the diagnosis is possible on the morphological features for adult worms for helminthologists, the identification on some stages (female, subadult, eggs and larvae) is difficult or impossible. Sequencing ND4 domain from the mitochondrial DNA of H. contortus and A. sidemi worms, we observed clearly two distinct clades, with an inter-specific divergence of 28.1%. Basing on this specific domain, a multiplex PCR-based method was developed: new primers were designed and used pooled in one mix PCR, producing amplicons of 454bp for H. contortus and 330bp for A. sidemi, allowing a trivial and an inexpensive taxonomic affiliation after migration. This multiplex PCR-based method was developed here to distinguish H. contortus and A. sidemi regardless their developmental stage, easy to use for highlighting co-infestation cases in both wild and domestic ruminants. It is a non-invasive approach appearing as a good diagnostic tool relevant to coprological cultures.

Guerrerostrongylus marginalis n. sp. (Trichostrongyloidea: Heligmonellidae) from the Guianan arboreal mouse (Oecomys auyantepui) from French Guiana.

Based on the number and arrangement of cuticular ridges and configuration of the dorsal ray, nematode specimens collected from the small intestine of eight Guianan arboreal mice, Oecomys auyantepui (Rodentia: Sigmodontinae), in French Guiana are herein described and characterized. Guerrerostrongylus marginalis n. sp. (Heligmosomoidea: Heligmonellidae) shows a synlophe consisting of more than 40 ridges and a unique bursal arrangement with ray 8 (externo-dorsal) extending to the edge of the bursal margin, and appearing more prominent than the dorsal ray. This bursal arrangement is common in members of Hassalstrongylus Durette-Desset, 1971, but uncommon in the other four species in Guerrerostrongylus Sutton & Durette-Desset, 1991. The placement of the new species in Guerrerostrongylus is based on the number and nature of cuticular ridges and the ray arrangement and symmetry of the caudal bursa. Diagnostic characteristics of Guerrerostrongylus marginalis n. sp. include the length of ray 8 relative to bursal margin, the relative size of the spicules and vestibule, and the number of eggs in the uterus. We propose an amendment to the generic diagnosis of Guerrerostrongylus to modify the characters of the long rays 6 (postero-lateral), rays 8 (externo-dorsal), and dorsal ray as diagnostic, since at least ray 6 appears to be short in two different species in the genus, namely G. ulysi Digiani, Notarnicola & Navone, 2012 and G. marginalis n. sp.

Insight into species diversity of the Trichostrongylidae Leiper, 1912 (Nematoda: Strongylida) in ruminants.

This paper focuses on the species diversity among the Trichostrongylidae Leiper, 1912 (Nematoda: Strongylida), and complexity of the family systematics. Polymorphism (subfamilies: Ostertagiinae, Cooperiinae and Haemonchinae), the presence of cryptic species (genus: Teladorsagia) and hybridization (genera: Cooperia, Haemonchus and Ostertagia) are presented and discussed, considering both morphological and molecular evidence. Some of these phenomena are common, nevertheless not sufficiently understood, which indicates the need for expanding the current state of knowledge thereof. Within the Trichostrongylidae, species distinction supported merely by morphological features is difficult, and requires confirmation by means of molecular methods. The parasitic nematode taxonomy is complicated mainly by the genus Teladorsagia, but complexity may also be expected among other Ostertagiinae (e.g. in the genera Ostertagia and Marshallagia). The data presented here show that the members of the Trichostrongylidae can significantly complicate unambiguous species identification. Hence, it is essential to consider the phenomena mentioned, to gather valid and comparable data on the biodiversity of this family.

Taxonomic revision of the Nippostrongylinae (Nematoda, Heligmonellidae) parasites of Muridae from the Australasian region. The genus Odilia Durette-Desset, 1973.

The species of the genus Odilia Durette-Desset, 1973 (Heligmonellidae, Nippostrongylinae) are re-distributed among eight genera of which five are new. This classification is mainly based on certain characters of the synlophe not previously taken into account at the supraspecific level. These characters mainly include the presence or absence of a careen, the relative size of the ridges forming the careen, the development and position of ridge 1', the development of the left ridge and right ridge, and the distribution of the largest ridges. Eighteen of the 20 known species are rearranged in the following genera: Odilia sensu stricto Durette-Desset, 1973 with Odilia mackerrasae (Mawson, 1961) as type species, Chisholmia n. gen. with Chisholmia bainae (Beveridge & Durette-Desset, 1992) n. comb. as type species, Equilophos n. gen. with Equilophos polyrhabdote (Mawson, 1961) n. comb. as type species, Hasegawanema n. gen. with Hasegawanema mamasaense (Hasegawa, Miyata & Syafruddin, 1999) n. comb. as type species, Hughjonestrongylus Digiani & Durette-Desset, 2014 with Hughjonestrongylus ennisae (Smales & Heinrich, 2010) as type species, Lesleyella n. gen. with Lesleyella wauensis (Smales, 2010) n. comb. as type and sole species, Parasabanema szalayi Smales & Heinrich, 2010, and Sanduanensis n. gen. with Sanduanensis dividua (Smales, 2010) as type and sole species. Odilia uromyos Mawson, 1961 and Odilia carinatae Smales, 2008 are not included in the new classification. A key to the proposed genera is provided. The new generic arrangement follows a distribution more related to the biogeographical areas than to the host groups.

A new species of Torrestrongylus (Trichostrongylidae, Anoplostrongylinae) from Macrotus waterhousii (Chiroptera: Phyllostomidae) in Central Mexico.

A new species of nematode, Torrestrongylus tetradorsalis n. sp., is described herein, based on specimens recovered from the small intestine of the leaf-nosed bat, Macrotus waterhousii, from the Biosphere Reserve "Sierra de Huautla" in the state of Morelos, Mexico. The new species is included in Torrestrongylus because it features a bursa of the type 3 - 2, a divided cephalic vesicle with an anterior half in the shape of an umbrella, and a posterior widened half. The new species can be distinguished from the only other congener T. torrei Pérez-Vigueras, 1935 by four key features: first, by the absence of cervical alae in both males and females; second, by the relatively longer second half of the cephalic cap; third, by the configuration of the dorsal ray, that does not have a medial terminal ray, and finally, by the structure of the spicules. This is the second species in the genus, previously known from bats of the families Phyllostomidae and Molossidae in Cuba, and now in Mexico.

Rediscovery and New Morphological Data on Two Hassalstrongylus (Nematoda: Heligmonellidae) Coparasitic in the Marsh Rat Holochilus chacarius (Rodentia: Cricetidae) from Argentina.

Two species of Hassalstrongylus Durette-Desset, 1971, coparasitic in Holochilus chacarius Thomas (Rodentia, Cricetidae) and not recorded since their original description in 1937, were newly found in their type host and locality. Hassalstrongylus mazzai (Freitas, Lent and Almeida, 1937) and Hassalstrongylus argentinus (Freitas, Lent and Almeida, 1937) were obtained from Ho. chacarius from 2 different populations: one from Salta Province (northwest Argentina) and another from Chaco Province (northeast Argentina). The species described as Heligmonoides mazzai Freitas, Lent and Almeida, 1937 had been transferred to Hassalstrongylus even though its synlophe had never been studied. We provide the first descriptions and illustrations of the synlophe of males and females of Hassalstrongylus mazzai and the female of H. argentinus and account for morphological and metrical variability. We confirm, through the study of the synlophe, the placement of Hassalstrongylus mazzai in the genus Hassalstrongylus and designate neotypes for the species because the type material deposited by the authors could not be found. Females of both species were morphologically very similar, and a principal components analysis (PCA) performed on some morphometrical characters showed that the body length, uterus length, and an unexpected character as the number of eggs were useful characters in the discrimination of both species.

The Status of Heligmosomoides americanus, Representative of an American Clade of Vole-Infecting Nematodes.

Heligmosomoides americanus is shown by molecular phylogenetic analysis of 3 nuclear (28S, ITS1, and ITS2) and 2 mitochondrial (cytochrome oxidase 1 and cytochrome b) loci to be a distinct species of heligmosomid nematode with a long-independent evolutionary history, and not a subspecies of Heligmosomoides polygyrus . Rather than being a recent arrival in North America, the species probably originated as a Beringian immigrant with the host vole Phenacomys, approximately 2 million years ago (MYA).

Comparative morphology of the species of Libyostrongylus and Codiostomum, parasites from ostriches, Struthio camelus, with a identification key to the species.

One of the most common problems in breeding of ostriches in captivity is the control of parasitic diseases. This work presents keys for the identification of adult nematodes and infective larvae by morphologic and morphometric characteristics. These keys will allow the scientific community to identify the species that infect the ostriches either based on the characteristics of the posterior end of the infective larvae found through a simple fecal exam or by observing the morphology and morphometry of adult worms recovered during necropsies. These keys will facilitate ecological and systematic studies, as well as increase the understanding of the epidemiology of these parasitosis in ostriches.

Comparative morphology of the species of Libyostrongylus and Codiostomum, parasites from ostriches, Struthio camelus, with a identification key to the species / Morfologia Comparativa das espécies de Libyostrongylus e Codiostomum, parasitas de avestruzes, Struthio camelus, com uma chave para identificação das espécies

One of the most common problems in breeding of ostriches in captivity is the control of parasitic diseases. This work presents keys for the identification of adult nematodes and infective larvae by morphologic and morphometric characteristics. These keys will allow the scientific community to identify the species that infect the ostriches either based on the characteristics of the posterior end of the infective larvae found through a simple fecal exam or by observing the morphology and morphometry of adult worms recovered during necropsies. These keys will facilitate ecological and systematic studies, as well as increase the understanding of the epidemiology of these parasitosis in ostriches.

Um dos problemas mais comuns na criação de avestruzes em cativeiro é o controle das doenças parasitárias. Este trabalho apresenta chaves para a identificação de Nematoda adultos e larvas infectantes através de caracteres morfológicos e morfométricos. Essas chaves de identificação permitirão à comunidade científica o diagnóstico das espécies que infectam as avestruzes com base nas características da extremidade posterior das larvas infectantes encontradas por meio de simples exames fecais ou pela observação da morfologia e morfometria dos espécimes adultos recuperados durante necropsia. Dessa forma, as chaves de identificação facilitarão os estudos ecológicos e sistemáticos, bem como a melhor compreensão da epidemiologia dessas infecções em avestruzes.

Population dynamics of Libyostrongylus dentatus and L. douglassii of ostriches raised in the Northern Rio de Janeiro State, Brazil.

Libyostrongylus genus contains three species of gastrointestinal nematodes that infect ostriches. Of these, only Libyostrongylus douglassii has been implicated in diseases and lower productivity. A morphological diagnosis method allowing the discrimination of infective larvae of L. douglassii and Libyostrongylus dentatus and the evaluation of eggs per gram of feces (EPG) was used to determine the population dynamics of both species from two ostrich farms located in the northern region of the state of Rio de Janeiro, Brazil. The presence of these nematodes was monitored every 15 days for two years. The average EPG was 4631 and 587 on farms 1 and 2, respectively. Overall, the highest values of EPG were associated with the rainy season on both farms. However, only on farm 1 the rainy season correlated statistically with EPG. Both species of Libyostrongylus occurred on both farms with a higher predominance (86%) of L. douglassii on farm 1, whereas the distribution of infection on farm 2 was more heterogeneous with a slight predominance of L. douglassii. The infection and the prevalence of these species may vary considerably according to the farm facilities, management adopted, and season.

Additional data on the synlophe of Stilestrongylus aculeata (Travassos, 1918) and Stilestrongylus eta (Travassos, 1937) (Heligmonellidae) parasitic in Akodon montensis (Sigmodontinae) from the Atlantic Forest.

Additional data on the synlophe and morphological measurements are provided for the heligmonellids Stilestrongylus aculeata (Travassos, 1918) and Stilestrongylus eta (Travassos, 1937), collected from Akodon montensis Thomas, 1913, of the Atlantic Forest, Rio de Janeiro, Brazil. The synlophe of S. aculeata is characterized by 19 cuticular middle body ridges in males and 24 cuticular middle body ridges in females, whereas S. eta presents 17 middle body ridges in males and 20 middle body ridges in females.

Molecular genetic conspecificity of Spiculopteragia houdemeri (Schwartz, 1926) and S. andreevae (Drózdz, 1965) (Nematoda: Ostertagiinae) from wild ruminants in Japan.

Male dimorphism of the subfamily Ostertagiinae (Nematoda: Trichostrongylidae) is a well-known phenomenon, and two or more morphotypes of a single species have previously been described as different species. Two Spiculopteragia spp., S. houdemeri (syn. S. yamashitai) and S. andreevae (syn. Rinadia andreevae) recorded in Asian cervids and wild bovids, are considered to represent major and minor morphs of S. houdemeri, respectively, based solely on their co-occurrence in the same host individual along with monomorphic females. In this study, males of morph houdemeri ( = S. houdemeri) and morph andreevae ( = S. andreevae) as well as females with three different vulval ornamentations were collected from sika deer (Cervus nippon) and Japanese serows (Capricornis crispus) distributed on the mainland of Japan. Morphologically characterized worms were subjected to molecular genetic analyses based on the internal transcribed spacer region of the ribosomal RNA gene and a partial region of the cytochrome c oxidase subunit I gene of mitochondrial DNA. Of 181 collected sika deer, 177 (97.8%) and 73 (40.3%) deer harboured males of morphs houdemeri and andreevae, respectively. Worm numbers of the former morph were found to range between 1 and 444 per individual, whereas only 1-25 worms per individual were detected for the latter morph. Five out of six serows harboured 47-71 or 2-9 males of morphs houdemeri and andreevae per individual, respectively. Females with one or two vulval flaps were predominant, but there was a substantial presence of flapless females in both host species. All the morphs of male and female adults had an identical genetic background, thus directly confirming the morphological polymorphism of S. houdemeri.