Molecular and morphological evidences reveal a cryptic species in the Vinaceous Rosefinch Carpodacus vinaceus (Fringillidae; Aves)

Wu, H.‐C., Lin, R.‐C., Hung, H.‐Y., Yeh, C.‐F., Chu, J.‐H., Yang, X.‐J., Yao, C.‐J., Zou, F.‐S., Yao, C.‐T., Li, S.‐H. & Lei, F.‐M. (2011). Molecular and morphological evidences reveal a cryptic species in the Vinaceous Rosefinch Carpodacus vinaceus (Fringillidae; Aves). —Zoologica Scripta, 40, 468–478. The Vinaceous Rosefinch (Carpodacus vinaceus) is endemic in East Asia with two recognized subspecies –C. v. vinaceus, distributed along the eastern edge of the Tibetan Plateau and the Himalayas, and C. v. formosanus, restricted to Taiwan’s Central Mountain Range. As reflected in a controversial taxonomic history, this vastly disjunctive distribution pattern suggests that the subspecies, having been isolated from each other for a long time, might have diverged, challenging the current taxonomic treatment and calling for possible species delimitation. Sequences of two mitochondrial fragments (mtDNA) and two Z‐linked nuclear loci (zDNA) were used to reconstruct the intraspecific phylogeny of C. vinaceous. The mtDNA tree shows that the two subspecies of the vinaceous rosefinch form two exclusively monophyletic clades. All but one zDNA sequences from the nominate subspecies and C. v. formosanus also formed exclusively monophyletic clades (the exceptional zDNA sequence from C. v. vinaceous formed a weakly supported clade with two outgroup species). Moreover, by conducting quantitative comparisons of morphometric traits and male plumage coloration, we found that the two subspecies exhibit distinguishable morphological differences. All the evidence therefore suggests that C. v. formosanus is a cryptic species and that its taxonomic status should be restored to full species. Molecular dating suggests that the two sibling rosefinches split 1.7 ± 0.2 million years ago, providing a point estimate for the historical connectivity of biota between eastern Tibet‐Himalayas and montane Taiwan.     

Morphological convergence in a Mexican garter snake associated with the ingestion of a novel prey

Morphological convergence is expected when organisms which differ in phenotype experience similar functional demands, which lead to similar associations between resource utilization and performance. To consume prey with hard exoskeletons, snakes require either specialized head morphology, or to deal with them when they are vulnerable, for example, during molting. Such attributes may in turn reduce the efficiency with which they prey on soft‐bodied, slippery animals such as fish. Snakes which consume a range of prey may present intermediate morphology, such as that of Thamnophiine (Natricinae), which may be classified morphometrically across the soft–hard prey dietary boundary. In this study, we compared the dentition and head structure of populations of Thamnophis melanogaster that have entered the arthropod–crustacean (crayfish)‐eating niche and those that have not, and tested for convergence between the former and two distantly related crayfish specialists of the genus Regina (R. septemvittata and R. grahamii). As a control, we included the congener T. eques. Multivariate analysis of jaw length, head length, head width, and number of maxillary teeth yielded three significant canonical variables that together explained 98.8% of the variance in the size‐corrected morphological data. The first canonical variable significantly discriminated between the three species. The results show that head dimensions and number of teeth of the two Regina species are more similar to those of crayfish‐eating T. melanogaster than to non‐crayfish‐eating snakes or of T. eques. It is unclear how particular head proportions or teeth number facilitates capture of crayfish, but our results and the rarity of soft crayfish ingestion by T. melanogaster may reflect the novelty of this niche expansion, and are consistent with the hypothesis that some populations of T. melanogaster have converged in their head morphology with the two soft crayfish‐eating Regina species, although we cannot rule out the possibility of a morphological pre‐adaptation to ingest crayfish.     

Parasite spillover: indirect effects of invasive Burmese pythons

Identification of the origin of parasites of nonindigenous species (NIS) can be complex. NIS may introduce parasites from their native range and acquire parasites from within their invaded range. Determination of whether parasites are non‐native or native can be complicated when parasite genera occur within both the NIS’ native range and its introduced range. We explored potential for spillover and spillback of lung parasites infecting Burmese pythons (Python bivittatus) in their invasive range (Florida). We collected 498 indigenous snakes of 26 species and 805 Burmese pythons during 2004–2016 and examined them for lung parasites. We used morphology to identify three genera of pentastome parasites, Raillietiella, a cosmopolitan form, and Porocephalus and Kiricephalus, both New World forms. We sequenced these parasites at one mitochondrial and one nuclear locus and showed that each genus is represented by a single species, R. orientalis, P. crotali, and K. coarctatus. Pythons are host to R. orientalis and P. crotali, but not K. coarctatus; native snakes are host to all three species. Sequence data show that pythons introduced R. orientalis to North America, where this parasite now infects native snakes. Additionally, our data suggest that pythons are competent hosts to P. crotali, a widespread parasite native to North and South America that was previously hypothesized to infect only viperid snakes. Our results indicate invasive Burmese pythons have affected parasite‐host dynamics of native snakes in ways that are consistent with parasite spillover and demonstrate the potential for indirect effects during invasions. Additionally, we show that pythons have acquired a parasite native to their introduced range, which is the initial condition necessary for parasite spillback.     

Complete mitochondrial genomes of prasinophyte algae Pyramimonas parkeae and Cymbomonas tetramitiformis

Mitochondria are archetypal eukaryotic organelles that were acquired by endosymbiosis of an ancient species of alpha‐proteobacteria by the last eukaryotic common ancestor. The genetic information contained within the mitochondrial genome has been an important source of information for resolving relationships among eukaryotic taxa. In this study, we utilized mitochondrial and chloroplast genomes to explore relationships among prasinophytes. Prasinophytes are represented by diverse early‐diverging green algae whose physical structures and genomes have the potential to elucidate the traits of the last common ancestor of the Viridiplantae (or Chloroplastida). We constructed de novo mitochondrial genomes for two prasinophyte algal species, Pyramimonas parkeae and Cymbomonas tetramitiformis, representing the prasinophyte clade. Comparisons of genome structure and gene order between these species and to those of other prasinophytes revealed that the mitochondrial genomes of P. parkeae and C. tetramitiformis are more similar to each other than to other prasinophytes, consistent with other molecular inferences of the close relationship between these two species. Phylogenetic analyses using the inferred amino acid sequences of mitochondrial and chloroplast protein‐coding genes resolved a clade consisting of P. parkeae and C. tetramitiformis; and this group (representing the prasinophyte clade I) branched with the clade II, consistent with previous studies based on the use of nuclear gene markers.     

Integrative taxonomy provides evidence for the species status of the Ibero‐Maghrebian grass snake Natrix astreptophora

The grass snake (Natrix natrix) is Europe's most widely distributed and, in many regions, most common snake species, with many morphologically defined subspecies. Yet, the taxonomy of grass snakes is relatively little studied and recent work has shown major conflicts between morphologically defined subspecies and phylogeographical differentiation. Using external morphology, osteological characters, and information from 13 microsatellite loci and two mitochondrial markers, we examine differentiation of the subspecies N. n. astreptophora from the North African Maghreb region, the Iberian Peninsula and neighbouring France. According to previous studies, N. n. astreptophora corresponds to a deeply divergent mitochondrial clade and constitutes the sister taxon of all remaining grass snakes. In the French Pyrenees region, there is a contact zone of N. n. astreptophora with another subspecies, N. n. helvetica. Our analyses of microsatellites and mitochondrial DNA reveal that the distribution ranges of the two taxa abut there, but both hybridize only exceptionally. Even though many morphological characters are highly variable and homoplastic in grass snakes, N. n. astreptophora differs consistently from all other grass snakes by its reddish iris coloration and in having significantly fewer ventral scales and another skull morphology. Considering further the virtual absence of gene flow between N. n. astreptophora and N. n. helvetica, and acknowledging the morphological distinctiveness of N. n. astreptophora and its sister group relationship to all remaining subspecies of grass snakes, we conclude that Natrix astreptophora (Seoane, 1884) should be recognized as a distinct species. Further research is needed to explore whether N. astreptophora is polytypic because a single sample of N. astreptophora from Tunisia turned out to be genetically highly distinct from its European conspecifics.     

Distribution patterns of four Termitomyces species cultivated by a fungus-growing termite,Odontotermes formosanus,in Taiwan

Fungus-growing termites inoculate the obligate symbiotic fungus Termitomyces into the fungus comb in their colonies. In this study, Taiwanese Termitomyces species were determined by diagnostic PCR using the metagenome of the body of Odontotermes formosanus (the only host termite living in Taiwan) as a template. Phylogenetic analyses revealed that four different Termitomyces species are cultivated by O. formosanus in Taiwan. Three have previously been registered in the DNA database, but one was first recorded in Taiwan. Only Termitomyces sp. Type C was distributed in all areas investigated in Taiwan, whereas the other three species were distributed regionally. Field observations indicated that the flush period and the number of fruit bodies in each colony varied between species. The distribution patterns of Termitomyces spp. in Taiwan may be related to the Taiwanese climate and/or the fruiting pattern.     

Length–weight relationships of three fish species from Moyangjiang River,China

Length‐weight relationships were determined for three fish species [Tachysurus argentivittatus (Regan, 1905); Rhodeus spinalis Oshima, 1926; Liniparhomaloptera disparis (Lin, 1934)]. All fish sample were collected using gillnets (15 × 5 m, mesh‐size 5 mm) from Moyangjiang River, China. Samples were collected quarterly from July 2006 to August 2007. The parameter b of length‐weight relationship varied from 2.52 for Rhodeus spinalis to 3.27 forLiniparhomaloptera disparis.     

Two sympatric new species of woodlizards (Hoplocercinae, Enyalioides) from Cordillera Azul National Park in northeastern Peru

We report the discovery of two sympatric new species of Enyalioides from a montane rainforest of the Río Huallaga basin in northeastern Peru. Among other characters, the first new species is distinguishable from other Enyalioides by the combination of the following characters: strongly keeled ventral scales, more than 37 longitudinal rows of dorsals in a transverse line between the dorsolateral crests at midbody, low vertebral crest on the neck with vertebrals on neck similar in size to those between hind limbs, projecting scales on body or limbs absent, 96 mm maximum SVL in both sexes, and caudals increasing in size posteriorly within each autotomic segment. The second new species differs from other species of Enyalioides in having strongly keeled ventral scales, scales posterior to the superciliaries forming a longitudinal row of strongly projecting scales across the lateral edge of the skull roof in adults of both sexes, 31 or fewer longitudinal rows of strongly keeled dorsals in a transverse line between the dorsolateral crests at midbody, vertebrals on neck more than five times the size of vertebrals between hind limbs in adult males, projecting scales on body or limbs absent, and caudals increasing in size posteriorly within each autotomic segment. We also present an updated molecular phylogenetic tree of hoplocercines including new samples of Enyalioides rudolfarndti, Enyalioides rubrigularis, both species described in this paper, as well as an updated identification key for species of Hoplocercinae.     

越南北部齿蛉及鱼蛉种类纪要（广翅目：齿蛉科）

基于近3年意大利佛罗伦萨大学与越南国家自然博物馆联合考察采集的标本,记述越南北部齿蛉科4属18种,包括6个越南新记录种:普通齿蛉Neoneuromus ignobilis Navás,1932、东方齿蛉N.orientalis Liu&Yang,2004、锡金齿蛉N.sikkimensis(van der Weele,1907)、广西星齿蛉Protohermesguangxiensis Yang&Yang,1986、台湾斑鱼蛉Neochauliodes formosanus(Okamoto,1910)和污翅斑鱼蛉N.fraternus(McLachlan,1869)。     

Description of a new genus with four new species of termite‐guest rove beetles (Coleoptera: Staphylinidae: Aleocharinae) associated with Hodotermopsis sjostedti (Isoptera: Termopsidae)

Hodotermopsiphila gen. nov. is described to include four termitophilous species: H. maruyamai sp. nov. (Yaku‐shima, Japan), H. nitens sp. nov. (Yaku‐shima, Japan), H. longisetosa sp. nov. (Taiwan), and H. sugayai sp. nov. (Taiwan and northern Laos). These species are associated with the damp‐wood termite, Hodotermopsis sjostedti Holmgren, 1911. Hodotermopsiphila species share possible apomorphic character states with three termitophilous species, Hodotermophilus gloriosus, Yakuus iwatai, and Termophidoholus formosanus, which are also associated with H. sjostedti.     

Phylogeography and diversification of an Amazonian understorey hummingbird: paraphyly and evidence for widespread cryptic speciation in the Plio‐Pleistocene

Straight‐billed Hermit Phaethornis bourcieri inhabits the understorey of upland terra firme forest throughout most of the Amazon basin. Currently, two allopatric taxa regarded as subspecies are recognized: P. b. bourcieri and P. b. major. However, the validity, interspecific limits and evolutionary history of these taxa are not yet fully elucidated. We use molecular characters to propose a phylogenetic hypothesis for populations and taxa grouped under Phaethornis bourcieri. Our results showed that P. bourcieri is part of the ‘Ametrornis’ clade, along with Phaethornis philippii and Phaethornis koepckeae, and that the subspecies major is more closely related to the latter two species than to populations grouped under nominate bourcieri. Our phylogenetic hypotheses recovered three main reciprocally monophyletic clades under nominate bourcieri separated by the lower Negro River and the Branco River or the Branco–Negro interfluve (clades B and C) and the upper Amazon (Solimões) or lower Marañon/Ucayali Rivers (clades C and D). Based on multi‐locus phylogeographic and population genetics approaches, we show that P. b. major is best treated as a separate species, and that P. b. bourcieri probably includes more than one evolutionary species, whose limits remain uncertain. The diversification of the ‘Ametrornis’ clade (P. bourcieri, P. philippii and P. koepckeae) is centred in the Amazon and appears to be closely linked to the formation of the modern Amazon drainage during the Plio‐Pleistocene.     

A time‐calibrated phylogenetic approach to assessing the phylogeography,colonization history and phenotypic evolution of snakes in the Japanese Izu Islands

Aim We infer the biogeography and colonization history of a dispersal‐limited terrestrial vertebrate, the Japanese four‐lined ratsnake (Elaphe quadrivirgata), to reveal the number of times mainland populations have invaded the Izu Archipelago of Japan, the mainland sources of these colonists, and the time‐scale of colonization. We compare these results with those of past studies in an attempt to uncover general biogeographical patterns. Moreover, we briefly examine the significance of colonization history when evaluating the evolution of body size and melanism of the Izu Island E. quadrivirgata populations. Location The Izu Islands (Oshima, Toshima, Niijima, Shikine, Kozu, Tadanae and Mikura), a volcanic archipelago off the Pacific coast of central Japan. Methods We obtained DNA sequences of the mitochondrial cytochrome b gene (1117 base pairs) from 373 individual snakes sampled from seven of the Izu Islands and 25 mainland localities. We employed partitioned Bayesian phylogenetic analyses assuming a relaxed molecular clock to estimate phylogenetic relationships among extant haplotypes and to give an explicit temporal scale to the timing of clade divergence, colonization history and tempo of body‐size evolution. Moreover, we employed model‐based biogeographical analysis to calculate the minimum number of times E. quadrivirgata colonized the Izu Islands. Results We found evidence that three separate regions of the Izu Archipelago have been colonized independently from mainland ancestors within the past 0.58–0.20 Ma. The Izu Peninsula plus Oshima and Mikura were both colonized independently from lineages inhabiting eastern mainland Japan. The Toshima, Niijima, Shikine, Kozu and Tadanae populations all derive from a single colonization from western mainland Japan. Oshima has been subject to three or possibly four colonizations. Main conclusions These results support the hypothesis that the extreme body‐size disparity among island populations of this ratsnake evolved in situ. Moreover, the fact that the dwarf, melanistic population inhabiting Oshima descends from multiple mainland colonization events is evidence of an extremely strong natural selection pressure resulting in the rapid evolution of this unique morphology. These results contrast with theoretical predictions that natural selection pressures should play a decreased role on islands close to the mainland and/or subject to frequent or recent immigration.     

Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) is not invasive through Asia: It's been there all along

Oriental fruit fly, Bactrocera dorsalis (Hendel), is a highly polyphagous fruit fly which, in the last 15 years, has invaded (with or without establishment) Africa, Europe and North America. As a direct result of these invasions, there is increasing research interest in the invasion history and spread patterns of this fly. A statement being repeatedly used in the B. dorsalis invasion literature is that the species was first identified from Taiwan in 1912 and that it subsequently spread through South‐East and South Asia during the 20th century. This assumption is incorrect and stems from: (a) an incomplete knowledge of B. dorsalis taxonomic history; and (b) a confounding of first taxonomic record with first invasion record. Rather than being first detected in Taiwan in 1912, the first record of oriental fruit fly was from “East India” (India orientali) under the synonymous name of Musca ferruginea by Fabricius in 1794, and in the 1910s, it was known not only from Taiwan, but widely across tropical Asia with records from India, Burma, Bengal, Sri Lanka (as Ceylon), Singapore and Indonesia (multiple islands). The taxonomic literature is very clear that oriental fruit fly has not invaded the rest of Asia from Taiwan since 1912, and this error should not continue to be repeated in the literature.     

Effects of dissociating agents on the fine structure of embryonic chick thyroid cells

Heads of the boid snakes Python sebae and Python molurus were dissected and the arthrology, myology and dentition studied. Living specimens of these species were observed and their feeding behavior analyzed by means of high- and regular-speed motion pictures. Camera speeds of up to 400 frames per second permitted examination of the jaw movements during the striking and seizing of prey. Motion picture studies conducted at regular speeds provided information on cranial movements during the swallowing of prey. The morphology of the head was correlated with observed movements in an attempt to analyze the functional and adaptive implications of the jaw apparatus. The cranial apparatus was discussed in terms of a linkage or kinematic chain whose constrainment and degrees of freedom were examined and compared with the jaw linkage of lizards. It was concluded that the very rigidly constrained mechanism in lizards is in remarkably sharp contrast to the very loose apparatus in snakes. Motions of various cranial bones were analyzed with particular attention given the mechanical factors involved. In full protraction the maxillae and palatines are lifted and rotated outward about a longitudinal axis. These movements are important in orienting the teeth with respect to the prey and are related to seizing and swallowing.     

TWO TYPES OF AUXILIARY CELL AMPULLAE IN GRATELOUPIA (HALYMENIACEAE,RHODOPHYTA), INCLUDING G. TAIWANENSIS SP. NOV. AND G. ORIENTALIS SP. NOV. FROM TAIWAN BASED ON rbcL GENE SEQUENCE ANALYSIS AND CYSTOCARP DEVELOPMENT1

Few species in the genus Grateloupia have been investigated in detail with respect to the development of the auxiliary cell ampullae before or after diploidization. In this study, we document the vegetative and reproductive structures of two new species of Grateloupia, G. taiwanensis S.‐M. Lin et H.‐Y. Liang sp. nov. and G. orientalis S.‐M. Lin et H.‐Y. Liang sp. nov., plus a third species, G. ramosissima Okamura, from Taiwan. Two distinct patterns are reported for the development of the auxiliary cell ampullae: (1) ampullae consisting of three orders of unbranched filaments that branch after diploidization of the auxiliary cell and form a pericarp together with the surrounding secondary medullary filaments (G. taiwanensis type), and (2) ampullae composed of only two orders of unbranched filaments in which only a few cells are incorporated into a basal fusion cell after diploization of the auxiliary cell and the pericarp consists almost entirely of secondary medullary filaments (G. orientalis type). G. orientalis is positioned in a large clade based on rbcL gene sequence analysis that includes the type species of Grateloupia C. Agardh 1822 , G. filicina. G. taiwanensis clusters with a clade that includes the generitype of Phyllymenia J. Agardh 1848 , Ph. belangeri from South Africa; that of Prionitis J. Agardh 1851 , Pr. lanceolata from Pacific North America; and that of Pachymeniopsis Y. Yamada ex Kawab. 1954, Pa. lanceolata from Japan. A reexamination of the type species of the genera Grateloupia, Phyllymenia, Prionitis, and Pachymeniopsis is required to clarify the generic and interspecific relationships among the species presently placed in Grateloupia.     

<Emphasis Type="Italic">Candida neustonensis</Emphasis> sp. nov., a novel ascomycetous yeast isolated from the sea surface microlayer in Taiwan

A new ascomycetous yeast species, Candida neustonensis is proposed in this study based on four strains (SN92^T, SN47, SJ22, SJ25) isolated from sea surface microlayer in Taiwan. These four yeast strains were morphologically, physiologically and phylogenetically identical to each other. No sexual reproduction was observed on 5% malt extract agar, corn meal agar, V8 agar, McClary’s acetate agar and potato-dextrose agar. Phylogenetic analysis of the sequences of the D1/D2 domain of the large subunit (LSU) rRNA gene places C. neustonensis as a member of the Pichia guilliermondii clade, it also reveals that the phylogenetically closest relatives of C. neustonensis are C. fukuyamaensis (4.4% divergence), C. xestobii (4.4% divergence) and P. guilliermondii (4.5% divergence). C. neustonensis also is clearly distinguished from other known species in the P. guilliermondii clade based on the results of physiology tests. From these comparison analyses, the following novel yeast species is proposed: Candida neustonensis sp. nov., with strain SN92^T (= BCRC 23108^T = JCM 14892^T = CBS 11061^T) as the type strain.     

Does maternal care evolve through egg recognition or directed territoriality?

The mechanism that facilitates the evolution of maternal care is ambiguous in egg‐laying terrestrial vertebrates: does the ability of mothers to recognize their own eggs lead them under some circumstances to begin providing care or can maternal care evolve from simply being in close proximity to the eggs (e.g. through territorial behaviour)? This question is difficult to answer because in most species, parental care is either absent altogether or present; in only a few species we have the opportunity to observe intraspecific variation in the expression of parental care. We studied a population of long‐tailed skinks (Eutropis longicaudata) in which females have recently evolved maternal care from a noncaring state. Females on Orchid Island, Taiwan, remain with their eggs during incubation and when doing so, actively deter egg predation by egg‐eating snakes (Oligodon formosanus); in all other populations, females lack post‐ovipositional maternal care. Nest‐guarding females on Orchid Island (i) showed antipredator behaviours only in the original nest site in which they laid eggs, even after we removed all of the eggs or substituted them with those of a conspecific; (ii) protect any eggs present inside the original nest site (even when the eggs belong to a conspecific); and (iii) develop this behaviour while gravid (i.e. prior to laying eggs). This supports the hypothesis that long‐tailed skinks cannot recognize their own eggs, suggesting that maternal care is a directed form of territoriality only expressed towards egg‐eating snakes and only during reproduction. Nest guarding is among the most primitive forms of parental care, and the recent evolution of this behaviour in a single population provides insight into one of the mechanisms by which parental care can originate in terrestrial vertebrates.     

Use of field‐portable ultrasonography reveals differences in developmental phenology and maternal egg provisioning in two sympatric viviparous snakes

A thorough understanding of the life cycles underlying the demography of wild species is limited by the difficulty of observing hidden life‐history traits, such as embryonic development. Major aspects of embryonic development, such as the rate and timing of development, and maternal–fetal interactions can be critical features of early‐life fitness and may impact population trends via effects on individual survival. While information on development in wild snakes and lizards is particularly limited, the repeated evolution of viviparity and diversity of reproductive mode in this clade make it a valuable subject of study. We used field‐portable ultrasonography to investigate embryonic development in two sympatric garter snake species, Thamnophis sirtalis and Thamnophis elegans in the Sierra Nevada mountains of California. This approach allowed us to examine previously hidden reproductive traits including the timing and annual variation in development and differences in parental investment in young. Both species are viviparous, occupy similar ecological niches, and experience the same annual environmental conditions. We found that T. sirtalis embryos were more developmentally advanced than T. elegans embryos during June of three consecutive years. We also found that eggs increased in volume more substantially across developmental stages in T. elegans than in T. sirtalis, indicating differences in maternal provisioning of embryos via placental transfer of water. These findings shed light on interspecific differences in parental investment and timing of development within the same environmental context and demonstrate the value of field ultrasonography for pursuing questions relating to the evolution of reproductive modes, and the ecology of development.     

Systematics of rove beetles (Coleoptera: Staphylinidae: Aleocharinae) associated with Hodotermopsis sjostedti (Isoptera: Termopsidae)

Members of the termitophilous subtribe Termitozyrina (Aleocharinae: Lomechusini) associated with Hodotermopsis sjostedti are taxonomically treated. The genera Hodotermophilus Naomi & Terayama and Termophidoholus Naomi & Hirono, and each type species (monotypic) are redescribed. Termophidoholus formosanus, originally described from Taiwan, and its host termite H. sjostedti are newly recorded from Laos. Yakuus iwatai Kanao & Maruyama gen. & sp. nov. and H. gloriosus were collected sympatrically in the same nests in Yaku‐shima, Japan. All of the above species, belonging to the three genera, share the presence of a batch of spurs at the tibial apex of fore and mid legs. Habitus photographs and illustrations of diagnostic features are provided for these species, and their phylogenetic relationships are discussed based on morphological similarities and the extant distribution of the host termite species.