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Birds are characterized by pneumatization of their skeletons by epithelial diverticula from larger, air-filled cavities. The diverticula-or 'air sacs'-that invade the postcranium result from outgrowths of the lungs; poslcranial pneumaticity has been very well studied. Much more poorly understood are the air sacs that pneumatize the skull. Study or craniofacial pneumaticity in modern birds (Neornithes) indicates the presence of two separate systems: nasal pneumaticity and tympanic pneumaticity, The lacrimal and maxillary bones arc pneumatized by diverticula of the main paranasal cavity, the antorbital sinus. There are five tympanic diverticula in neornithines that pneumatize the quadrate, articulare and the bones of the braincase. The pneumatic features of the following six genera of Mesozoic birds are examined: Archaeopteryx, Ellaliornis, Baplomis, Parahesperornis, Hesperornis and lchthyornis. Despite the 'archaic' aspect of most of these birds, many of the pneumatic features of neornithines are found in .Mesozoic birds and are considered primitive for Aves. The phylogenetic levels at which most of the avian pneumatic features arose within Archosauria are uncertain. Until the phylogenetic levels at which homologous pneumatic features arose are determined, it is unwise to use most pneumatic characters in the discussion of avian origins. Within avian phylogeny, Ornithurae and Neornithes are well-supported by pneumatic synapomorphies. There is a trend towards reduction of craniofacial pneumaticity within Hesperornithiformes. Witthin Neornithes, four derived pneumatic characters suggest that the Palaeognathae (ratites and tinamous) is monophyletic.
A cladistic analysis is presented of the hawkmoths of the tribe Acherontiini, Morgan´s Sphinx (Xanthopan morganii (Walker», and related genera. The study aims to test the monophyly of tribe Acherontiini; the hypothesis that all taxa with extremely long probosces (some Acherontiini, Meganoton rubescens, Neococytius, Xanthopan) form a monophyletic group, or at least fall within a single reasonably compact clade; and, within this group, to determine whether Xanthopan is more closely related to Acherontiini or to COCytillS and Neococytius. The data set comprises 109 characters derived from adult and immature stage morphology, biology and behaviour. These data were analysed using equal weighting, successive approximations character weighting (SACW) and implied weighting. All weighting schemes agreed on the monophyly of Acherontiini and of a group of genera comprising Amphimoea, Cocytius and Neococytius (the Cocytius group). Several other generic and suprageneric clades were also consistently recovered. However, those hawkmoths with extremely long probosces were never recovered as a monophyletic group. The relationships of Xanthopan were also ambiguous. Equal weighting and SACW placedXanthopan + Meganoton rztbescens (Butler) as sister to the COCytills group, while implied weighting placed Xanthopan as sister to Acherontiini. This latter relationship is based primarily on shared possession of a pilifer/palp hearing organ. Further analyses suggested the two components of this organ were not biologically independent. Downweighting this feature accordingly resulted in all weighting schemes converging on the topology found by equal weighting. Exclusion of the incomplete subset of immature stage data had no effect under implied weighting but equal weighting and SACW now recovered a Neotropical clade comprising Manduca. and the Cocytius group, while Xanthopan was placed with M. rubescens and Panogena. Downweighting the pilifer/palp hearing organ under implied weighting again caused convergence with the equal weighting/SACW results. Thus, the relationships of Xanthopan remain equivocal and further data, particularly from the immature stages, will be required to elucidate its phylogenetic position further.
The taxonomy, diversity, and distribution of the aquatic insect order Trichoptera, caddisflies, are reviewed. The order is among the most important and diverse of all aquatic taxa. Larvae are vital participants in aquatic food webs and their presence and relative abundance are used in the biological assessment and monitoring of water quality. The species described by Linnaeus are listed. The morphology of all life history stages (adults, larvae, and pupae) is diagnosed and major features of the anatomy are illustrated. Major components of life history and biology are summarized. A discussion of phylogenetic studies within the order is presented, including higher classification of the suborders and superfamilies, based on recent literature. Synopses of each of 45 families are presented, including the taxonomic history of the family, a list of all known genera in each family, their general distribution and relative species diversity, and a short overview of family-level biological features. The order contains 600 genera, and approximately 13,000 species.
Lepidoptera phylogeny and systematics : the state of inventorying moth and butterfly diversity
(2007)
The currently recognized robust support for the monophyly of the Lepidoptera (and the superorder Amphiesmenoptera comprising Lepidoptera + Trichoptera) is outlined, and the phylogeny of the principal lineages within the order is reviewed succinctly. The state of the taxonomic inventory of Lepidoptera is discussed separately for ‘micro-moths’, ‘macro-moths’ and butterflies, three assemblages on which work has followed historically somewhat different paths. While currently there are about 160,000 described species of Lepidoptera, the total number of extant species is estimated to be around half a million. On average, just over one thousand new species of Lepidoptera have been described annually in recent years. Allowing for the new synonyms simultaneously established, the net increase in species numbers still exceeds 800/year. Most of the additions are foreseeable in the micro-moth grade, but even for butterflies ca 100 species are added annually. Examples of particularly interesting new high-rank taxa that have been described (or whose significance has become realized) since the middle of the 20th century include the non-glossatan lineages represented by Agathiphaga and Heterobathmia and the heteroneuran families Andesianidae, Palaephatidae, Hedylidae and Micronoctuidae. Some thoughts on how present and future systematic lepidopterology might be prioritised are presented.
Camel spiders (Arachnida: Solifugae) are one of the arachnid groups characterised by a prosomal dorsal shield composed of three distinct elements: the pro-, meso- and metapeltidium. These are associated respectively with prosomal appendages one to four, five, and six. What is less well known, although noted in the historical literature, is that the coxae of the 4th and 5th prosomal segments (i.e. walking legs 2 and 3) of camel spiders are also separated ventrally by a distinct membranous region, which is absent between the coxae of the other legs. We suggest that this essentially ventral division of the prosoma specifically between coxae 2 and 3 is homologous with the so-called sejugal furrow (the sejugal interval sensu van der Hammen). This division constitutes a fundamental part of the body plan in acariform mites (Arachnida: Acariformes). If homologous, this sejugal furrow could represent a further potential synapomorphy for (Solifugae + Acariformes); a relationship with increasing morphological and molecular support. Alternatively, outgroup comparison with sea spiders (Pycnogonida) and certain early Palaeozoic fossils could imply that the sejugal furrow defines an older tagma, derived from a more basal grade of organisation. In this scenario the (still) divided prosoma of acariform mites and camel spiders would be plesiomorphic. This interpretation challenges the textbook arachnid character of a peltidium (or ‘carapace’) covering an undivided prosoma.
The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38 tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample (37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on the morphological examination of many representatives of the subfamily, the classification of the tribes and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of the included genera, and notes on their distribution, ecology, and important vector species are provided for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported by a comprehensive database for Deltocephalinae compiled using the taxonomic database software package 3I. The online functionality includes an interactive key to tribes and subtribes and advanced database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage providing nomenclatural information, lists of included taxa, an automated description (if available), images (if available), distributional information, bibliographic references and links to outside resources. Some observations and trends regarding the history of taxonomic descriptions in Deltocephalinae are reported. Four new tribes are described: Bahitini tribe nov. (25 genera), Bonsapeiini tribe nov. (21 genera), Phlepsiini tribe nov. (4 genera), and Vartini tribe nov. (7 genera). The circumscription and morphological characterization of Scaphoideini Oman, 1943 (61 genera) is substantially revised. Eleven new species are described: Acostemma stilleri sp. nov., Arrugada linnavuorii sp. nov., Drabescus zhangi sp. nov., Parabolopona webbi sp. nov., Goniagnathus emeljanovi sp. nov., Hecalus hamiltoni sp. nov., Scaphoideus omani sp. nov., Dwightla delongi sp. nov., Abimwa knighti sp. nov., Gannia viraktamathi sp. nov., and Doratulina dmitrievi sp. nov. Some family-group level taxonomic changes are made: Platymetopiini Haupt, 1929, Anoterostemmini Haupt, 1929, and Allygidiina Dmitriev, 2006 are synonymized with Athysanini Van Duzee, 1892, syn. nov.; Procepitini Dmitriev, 2002 is synonymized with Cicadulini Van Duzee, 1892, syn. nov.; Listrophorini Boulard, 1971 is synonymized with Chiasmini Distant, 1908, syn. nov.; Adamini Linnavuori & Al-Ne’amy, 1983, Dwightlini McKamey, 2003, and Ianeirini Linnavuori, 1978 are synonymized with Selenocephalini Fieber, 1872 syn.nov., and all three are now recognized as valid subtribes in their parent tribe. New placements of many genera to tribe and subtribe are made, and these are described in individual taxon treatments.
Cteniogaster, a new genus of small ground spiders is described from Kenya and Tanzania. It encompasses seven new species, three of which are known from both sexes: C. toxarchus sp. nov., the type species, C. conviva sp. nov. and C. hexomma sp. nov. Three species are known from females only: C. lampropus sp. nov., C. sangarawe sp. nov. and C. taxorchis sp. nov. and one only from males: C. nana sp. nov. The new genus can be recognised by the presence of a posterior ventral abdominal f eld of strong setae and anterior lateral spinnerets with enlarged piriform gland spigots in males. A cladistic analysis attributes the genus to Liocranidae, Cybaeodinae. The results of the analysis performed do not produce an unequivocal autapomorphy for Liocranidae, but provide a combination of non-homoplasious character changes that offers significant potential for recognising genera as Liocranidae. Moreover, robust apomorphies are determined within Liocranidae for the subfamilies Liocraninae and Cybaeodinae. Based on these fi ndings Toxoniella Warui & Jocqué, 2002 is transferred from Gallieniellidae to Liocranidae, Cybaeodinae. Jacaena Thorell, 1897, Plynnon Deeleman-Reinhold, 2001 and Teutamus Thorell, 1890 are transferred to Corinnidae, Phrurolithinae and Montebello Hogg, 1914 to Gnaphosidae. Itatsina Kishida, 1930 is synonymised with Prochora Simon, 1886.
Background: The current taxonomy of the African giraffe (Giraffa camelopardalis) is primarily based on pelage pattern and geographic distribution, and nine subspecies are currently recognized. Although genetic studies have been conducted, their resolution is low, mainly due to limited sampling. Detailed knowledge about the genetic variation and phylogeography of the South African giraffe (G. c. giraffa) and the Angolan giraffe (G. c. angolensis) is lacking. We investigate genetic variation among giraffe matrilines by increased sampling, with a focus on giraffe key areas in southern Africa.
Results: The 1,562 nucleotides long mitochondrial DNA dataset (cytochrome b and partial control region) comprises 138 parsimony informative sites among 161 giraffe individuals from eight populations. We additionally included two okapis as an outgroup. The analyses of the maternally inherited sequences reveal a deep divergence between northern and southern giraffe populations in Africa, and a general pattern of distinct matrilineal clades corresponding to their geographic distribution. Divergence time estimates among giraffe populations place the deepest splits at several hundred thousand years ago.
Conclusions: Our increased sampling in southern Africa suggests that the distribution ranges of the Angolan and South African giraffe need to be redefined. Knowledge about the phylogeography and genetic variation of these two maternal lineages is crucial for the development of appropriate management strategies.
We compared Chatham Island endemic species Xanthocnemis tuanuii to its congenerics from the New Zealand South Island: X. zealandica (newly collected specimens)and X. sinclairi (type specimens plus newly collected material). Two independent tests were performed –geometric morphometrics and molecular. Both analyses were consistent in supporting the status of X. tuanuiias a good species. Species differed statistically in the following morphological traits: head (dorsal view), male appendages (dorsal, lateral, posterior and ventral views), thorax (dorsal view), and penis (dorsal and lateral view). In addition to the original diagnostic features (mainly shape of the male superior appendages), a new morphological character is suggested here which reliably distinguishes the species based on the shape of the inferior appendages. There was no statistical support for the species status of X. sinclairi. The only feature re-ported as diagnostic (lower lobe of male superior appendages) was found to be variable and insufficient to warrant the previously proposed taxonomic rank for X. sinclairi. Molecular analysis of specimens showing identical appendages to the X. sinclairi holotype grouped them with X. zealandica specimens. Therefore X. sinclairi is synonymised with X. zealandica.
This study deals with the biodiversity and distribution of cavernicolous Amphipoda in caves of the Arabika massif (Western Caucasus). The Sarma, Trojka and Orlinoe Gnezdo caves were explored during speleological expeditions over the years 2011–12. Two new species of Amphipoda were found: a sub-surface dweller Zenkevitchia sandroruffoi sp. nov. is reported from the Sarma, Trojka and Orlinoe Gnezdo caves at depths from -30 m to -350 m; the second one, a deep dweller Adaugammarus pilosus gen. et sp. nov. is reported from the Sarma Cave at depths of -1270 to -1700 m. Adaugammarus gen. nov. shares similarities with Typhlogammarus Schäferna, 1907 and Zenkevitchia Birstein, 1940. The species Anopogammarus birsteini Derzhavin, 1945 is also re-described herein based on new samples that suggest close affinity of this species with the family Gammaridae. The original taxonomic combination is resurrected for Zenkevitchia revazi Birstein & Ljovuschkin, 1970, comb. resurr. (from Anopogammarus Derzhavin, 1945). To accommodate morphologically different species in the genus Zenkevitchia, two new groups are proposed. These are the admirabilis-group (Z. admirabilis Birstein, 1940 and Z. yakovi Sidorov, 2015) and the sandroruffoi-group (Z. sandroruffoi sp. nov. and Z. revazi). An updated molecular (mt-cox1) phylogeny, an identification key to the genera and a distribution map for the typhlogammarid amphipod species of Transcaucasia are provided.
The Thyropygus opinatus subgroup (Diplopoda: Harpagophoridae) of the T. allevatus group in Thailand is revised. Based on a phylogenetic analysis of mtDNA sequence data, it is merged with the T. bifurcus subgroup to form an extended T. opinatus subgroup. Nine new species are described: Thyropygus cimi sp. nov. and T. forceps sp. nov. from Nakhonsrithammarat Province, T. culter sp. nov., T. planispina sp. nov., T. undulatus sp. nov. and T. ursus sp. nov. from Krabi Province, T. mesocristatus sp. nov. from Songkhla Province, T. navychula sp. nov. from Phang-Nga Province and T. sutchariti sp. nov. from Phetchaburi Province.
The New World genus Chariessa Forster (Coleoptera: Cleroidea: Cleridae) is revised and includes C. catalina Opitz, new species, C. elegans Horn, C. dichroa (LeConte), C. floridana Schaeffer, C. pilosa (Forster), C. texana Wolcott, C. ramicornis Perty, C. vestita (Chevrolat), and C. duponti (Spinola). Enoplium pilosa var. marginata Say is synonymized with Chariessa pilosa Forster. Lectotypes are designated for C. pilosa (Forster), C. ramicornis Perty, and C. vestita (Chevrolat). Available information indicates that Chariessa adult and immature individuals are predatory on lignicolous insects with a particular affinity for cerambycids and buprestids that infest species of oak. It is postulated that Pleistocene speciation generated the North American components of Chariessa with more ancient southern species generated during the Middle Tertiary; after closures of the Middle American portals and orogeny of the South American Andes. Included in this treatise is a discussion of natural history, key to species, narratives of zoogeography and phylogeny, one diagram of a phylogenetic tree, 35 line drawings, eight SEM micrographs, twelve habitus photographs, nine photographs of male genitalia, and five distributional maps.
Phylogenetic reconstruction from transposable elements (TEs) offers an additional perspective to study evolutionary processes. However, detecting phylogenetically informative TE insertions requires tedious experimental work, limiting the power of phylogenetic inference. Here, we analyzed the genomes of seven bear species using high-throughput sequencing data to detect thousands of TE insertions. The newly developed pipeline for TE detection called TeddyPi (TE detection and discovery for Phylogenetic Inference) identified 150,513 high-quality TE insertions in the genomes of ursine and tremarctine bears. By integrating different TE insertion callers and using a stringent filtering approach, the TeddyPi pipeline produced highly reliable TE insertion calls, which were confirmed by extensive in vitro validation experiments. Analysis of single nucleotide substitutions in the flanking regions of the TEs shows that these substitutions correlate with the phylogenetic signal from the TE insertions. Our phylogenomic analyses show that TEs are a major driver of genomic variation in bears and enabled phylogenetic reconstruction of a well-resolved species tree, despite strong signals for incomplete lineage sorting and introgression. The analyses show that the Asiatic black, sun, and sloth bear form a monophyletic clade, in which phylogenetic incongruence originates from incomplete lineage sorting. TeddyPi is open source and can be adapted to various TE and structural variation callers. The pipeline makes it possible to confidently extract thousands of TE insertions even from low-coverage genomes (∼10×) of nonmodel organisms. This opens new possibilities for biologists to study phylogenies and evolutionary processes as well as rates and patterns of (retro-)transposition and structural variation.
The Australian wolf spider genus Tetralycosa Roewer, 1960, with Lycosa meracula Simon, 1909 (junior synonym of Lycosa oraria L. Koch, 1877) as type species, is revised to include 13 species, eight of which are described as new here: Tetralycosa adarca sp. nov., T. alteripa (McKay, 1976), T. arabanae Framenau, Gotch & Austin, 2006, T. baudinettei sp. nov., T. caudex sp. nov., T. eyrei (Hickman, 1944), T. floundersi sp. nov., T. halophila sp. nov., T. oraria (L. Koch, 1876), T. orariola sp. nov., T. williamsi sp. nov., T. wundurra (McKay, 1979) comb. nov. and T. rebecca sp. nov. Members of Tetralycosa are halotolerant, exclusively inhabiting saline environments such as coastal beaches, and mound springs, clay pans and salt lakes in the Australian interior. A phylogenetic analysis of the genus identified a monophyletic clade of eight species that live permanently on the barren surface of salt lakes suggesting a single radiation into this extremely inhospitable habitat. Some of these Tetralycosa species are currently known from single salt lakes only and with increasing disturbances of these systems by mining, agriculture an
Acrodiscus Zanardini is a poorly known monotypic endemic Mediterranean genus based on A. vidovichii (Menegh.) Zanardini. Rarely reported, its reproductive structures have remained undocumented, leaving its exact taxonomic position uncertain. Solely on the basis of its vegetative structure, Zanardini provisionally placed it in the family Cryptonemiaceae of the order Cryptonemiales (currently the Halymeniaceae of the Halymeniales), although he was uncertain as to whether the new genus actually belonged to that family or should instead be included in the Gigartinaceae of the Gigartinales (where Meneghini had originally placed it). In the present study we have extensively sampled A. vidovichii and documented its vegetative and tetrasporangial features. As well, we provide molecularsequence data (COI-5P, rbcL, LSU) that indicate its phylogenetic affinities. We confirm Acrodiscus as a member of the Halymeniaceae and its status as an independent genus. Searches of several institutional herbaria have allowed us to locate and lectotypify Meneghini’s Chondrus? vidovichii by the discovery of his original material now held at the Herbarium Horti Pisani (Pisa, Italy).
With 280 accepted species, the genus Riccardia S.F.Gray (Aneuraceae) is one of the most speciose genera of simple thalloid liverworts. The current classification of this genus is based on morphological and limited-sampling molecular studies. Very few molecular data are available and a comprehensive view of evolutionary relationships within the genus is still lacking. A phylogeny focusing on relationships within the large genus Riccardia has not been conducted. Here we propose the first worldwide molecular phylogeny of the genus Riccardia, based on Bayesian inference and parsimony ratchet analyses of sequences from three plastid regions (psbA-trnH, rps4, trnL-F). The results support the monophyly of Riccardia and a new monospecific genus, Afroriccardia Reeb & Gradst. gen. nov., is described based on molecular and morphological evidence. The results indicate that several currently recognized infrageneric divisions and a few species are not monophyletic, suggesting that further analyses are needed to arrive at a proper understanding of the phylogeny of the genus. Although evidence for an Andean clade was found, most of the species appear scattered in different clades without clear geographical segregation. Broader sampling and further analyses are necessary in order to improve our understanding of the phylogeny of this poorly known liverwort genus.
A new classification of Ophiuroidea, considering family rank and above, is presented. The new family and superfamily taxa in O’Hara et al. (2017) were proposed to ensure a better readability of the new phylogeny but are unavailable under the provisions of the ICZN. Here, the morphological diagnoses to all 33 families and five superfamilies are provided. Ten new families, Ophiosphalmidae fam. nov., Ophiomusaidae fam. nov., Ophiocamacidae fam. nov., Ophiopteridae fam. nov., Clarkcomidae fam. nov., Ophiopezidae fam. nov., Ophiernidae fam. nov., Amphilimnidae fam. nov., Ophiothamnidae fam. nov. and Ophiopholidae fam. nov., are described. The family Ophiobyrsidae Matsumoto, 1915, not yet discovered in the previous publication, is added, based on new molecular data. A new phylogenetic reconstruction is presented. Definitions of difficult-to-apply morphological characters are given.
Synchroidae Lacordaire, 1859 is a taxonomically and biologically poorly known group. In the present paper, diagnostic characters used to separate genera are analysed and the phylogenetic relationships within this family are preliminarily investigated. Results suggest that the characteristic Synchroa pangu Hsiao, Li, Liu & Pang, 2016 can be removed to establish a new genus, Thescelosynchroa gen. nov. The new combination, T. pangu (Hsiao, Li, Liu & Pang) gen. et comb. nov., is proposed. The definitions of Synchroa Newman, 1838 and Synchroina Fairmaire, 1898 are revised. Moreover, morphological analysis and character comparison also suggest that the familial placement of Mallodrya subaenea Horn, 1888 is questionable. Six species are re-examined and rediagnosed: Synchroa chinensis Nikitsky, 1999, S. elongatula Nikitsky, 1999, S. formosana Hsiao, 2015, S. melanotoides Lewis, 1895, S. punctata Newman, 1838 and Synchroina tenuipennis Fairmaire, 1898. The male of S. chinensis and the female of S. formosana are described for the first time. Synchroa elongatula and Synchroina tenuipennis are newly recorded from Laos and Indonesia, respectively. We also hypothesize that the Eastern Asian-North American disjunction of Synchroa could be connected to a Mid-Late Tertiary migration of plants via the Bering Land Bridge.
In this paper we describe Macrobiotus canaricus sp. nov., a new tardigrade species of the Macrobiotus hufelandi group from the Canary Islands. Moreover, with the use of DNA sequencing, we confirm that Macrobiotus recens Cuénot, 1932 represents the hufelandi group, even though eggs laid by this species do not exhibit the typical hufelandi group morphology. Our study is based on both classical taxonomic methods that include morphological and morphometric analyses conducted with the use of light and scanning electron microscopy, and on the analysis of nucleotide sequences of four molecular markers (three nuclear: 18S rRNA, 28S rRNA, ITS-2, and one mitochondrial: COI). Our analyses revealed that M. canaricus sp. nov. is most similar to Macrobiotus almadai Fontoura et al., 2008 from the Archipelago of the Azores, from which it differs by the absence of granulation patches on the external and internal surfaces of legs I–III as well as by the absence of a cuticular pore in the centre of the external patch on legs I–III. Molecular sequences allowed us to pinpoint the phylogenetic positions of M. canaricus sp. nov. and M. recens within the M. hufelandi group.
Morphological and allozyme analyses suggested the occurrence of a pseudocryptic species in the Lasioglossum villosulum (Kirby, 1802) species complex (Hymenoptera: Halictidae). We analysed the morphology of more than 1500 specimens and the DNA barcode fragment of the cytochrome c oxidase subunit I (COI) of 102 specimens of this species complex from several Palaearctic countries. Our phylogenetic tree reconstructions, based on maximum likelihood and Bayesian inference revealed one clade corresponding to all specimens morphologically identified as Lasioglossum medinai (Vachal, 1895) and one divergent specimen morphologically identified as Lasioglossum berberum (Benoist, 1941). The other specimens, morphologically identified as L. villosulum, aggregated into at least three other lineages in our phylogenetic trees. The tree-based species delineations methods based on the Generalized Mixed Yule Coalescent (GMYC) model and the Bayesian Poisson Tree Process (bPTP) identified five to ten candidate species within the L. villosulum species complex, with L. medinai and L. berberum consistently recognized as separated from all other candidate species. Diagnostic morphological differences were found among L. medinai, L. berberum and the remaining specimens identified as L. villosulum. No diagnostic morphological differences were found to distinguish the different phylogenetic candidate species or lineages found within L. villosulum and L. medinai. Thus, both genetic and morphological approaches support the existence of L. medinai and L. berberum as distinct species from L. villosulum.